Adaptation to host plants may prevent rapid insect responses to climate change

We must consider the role of multitrophic interactions when examining species' responses to climate change. Many plant species, particularly trees, are limited in their ability to shift their geographic ranges quickly under climate change. Consequently, for herbivorous insects, geographic mosaics of host plant specialization could prohibit range shifts and adaptation when insects become separated from suitable host plants. In this study, we examined larval growth and survival of an oak specialist butterfly (Erynnis propertius) on different oaks (Quercus spp.) that occur across its range to determine if individuals can switch host plants if they move into new areas under climate change. Individuals from Oregon and northern California, USA that feed on Q. garryana and Q. kelloggii in the field experienced increased mortality on Q. agrifolia, a southern species with low nutrient content. In contrast, populations from southern California that normally feed on Q. agrifolia performed well on Q. agrifolia and Q. garryana and poorly on the northern, high elevation Q. kelloggii. Therefore, colonization of southern E. propertius in higher elevations and some northern locales may be prohibited under climate change but latitudinal shifts to Q. garryana may be possible. Where shifts are precluded due to maladaptation to hosts, populations may not accrue warm‐adapted genotypes. Our study suggests that, when interacting species experience asynchronous range shifts, historical local adaptation may preclude populations from colonizing new locales under climate change.     

Acclimation of photosynthetic temperature optima of temperate and boreal tree species in response to experimental forest warming

Rising temperatures caused by climate change could negatively alter plant ecosystems if temperatures exceed optimal temperatures for carbon gain. Such changes may threaten temperature‐sensitive species, causing local extinctions and range migrations. This study examined the optimal temperature of net photosynthesis (T_opt) of two boreal and four temperate deciduous tree species grown in the field in northern Minnesota, United States under two contrasting temperature regimes. We hypothesized that T_opt would be higher in temperate than co‐occurring boreal species, with temperate species exhibiting greater plasticity in T_opt, resulting in better acclimation to elevated temperatures. The chamberless experiment, located at two sites in both open and understory conditions, continuously warmed plants and soils during three growing seasons. Results show a modest, but significant shift in T_opt of 1.1 ± 0.21 °C on average for plants subjected to a mean 2.9 ± 0.01 °C warming during midday hours in summer, and shifts with warming were unrelated to species native ranges. The 1.1 °C shift in T_opt with 2.9 °C warming might be interpreted as suggesting limited capacity to shift temperature response functions to better match changes in temperature. However, T_opt of warmed plants was as well‐matched with prior midday temperatures as T_opt of plants in the ambient treatment, and T_opt in both treatments was at a level where realized photosynthesis was within 90–95% of maximum. These results suggest that seedlings of all species were close to optimizing photosynthetic temperature responses, and equally so in both temperature treatments. Our study suggests that temperate and boreal species have considerable capacity to match their photosynthetic temperature response functions to prevailing growing season temperatures that occur today and to those that will likely occur in the coming decades under climate change.     

Effects of rising temperatures and [CO2] on the physiology of tropical forest trees

Using a mixture of observations and climate model outputs and a simple parametrization of leaf-level photosynthesis incorporating known temperature sensitivities, we find no evidence for tropical forests currently existing "dangerously close" to their optimum temperature range. Our model suggests that although reductions in photosynthetic rate at leaf temperatures (TL) above 30 degrees C may occur, these are almost entirely accountable for in terms of reductions in stomatal conductance in response to higher leaf-to-air vapour pressure deficits D. This is as opposed to direct effects of TL on photosynthetic metabolism. We also find that increases in photosynthetic rates associated with increases in ambient [CO2] over forthcoming decades should more than offset any decline in photosynthetic productivity due to higher D or TL or increased autotrophic respiration rates as a consequence of higher tissue temperatures. We also find little direct evidence that tropical forests should not be able to respond to increases in [CO2] and argue that the magnitude and pattern of increases in forest dynamics across Amazonia observed over the last few decades are consistent with a [CO2]-induced stimulation of tree growth.     

Relationships between change in fire frequency and mortality due to spruce budworm outbreak in the southeastern Canadian boreal forest

Abstract. We present a simple empirical model that allows an estimation of mortality due to spruce budworm (Choristoneura fumiferana) outbreak in relation to fire frequency and site characteristics. The occurrence of a recent spruce budworm outbreak around Lake Duparquet (48° 30’N, 79° 20’W, ca. 300 m a.s.l.) in northwestern Québec permitted a reconstruction of the stand composition before the outbreak, and also of the mortality of Abies balsamea due to the outbreak. The basal area of A. balsamea increases with time since fire in all site types but with increasing values for (1) rock and shallow till, via (2) till and mesic clay up to (3) hydric clay. Mortality (measured as percentage loss of basal area due to the outbreak) increases with time since fire but did not vary with site type. The increasing abundance of A. balsamea with time since fire is mainly responsible for this increase in mortality. Mortality for a specific basal area is, however, lower for the more recently burned stands suggesting a significant residual effect of time since fire. A landscape model integrating mortality due to the outbreak for stands of different age is developed. Both absolute and relative losses of basal area increased with the length of the fire cycles. According to this model, changes in fire cycle could explain a large portion of the spatio-temporal variations observed in outbreak mortality in the southeastern boreal forest of Canada.     

Observed forest trait velocities have not kept pace with hydraulic stress from climate change

The extent to which future climate change will increase forest stress and the amount to which species and forest ecosystems can acclimate or adapt to increased stress is a major unknown. We used high-resolution maps of hydraulic traits representing the diversity in tree drought tolerance across the United States, a hydraulically enabled tree model, and forest inventory observations of demographic shifts to quantify the ability for within-species acclimation and between-species range shifts to mediate climate stress. We found that forests are likely to experience increases in both acute and chronic hydraulic stress with climate change. Based on current species distributions, regional hydraulic trait diversity was sufficient to buffer against increased stress in 88% of forested areas. However, observed trait velocities in 81% of forested areas are not keeping up with the rate required to ameliorate projected future stress without leaf area acclimation.     

Interactive climate factors restrict future increases in spring productivity of temperate and boreal trees

Climate warming is currently advancing spring leaf‐out of temperate and boreal trees, enhancing net primary productivity (NPP) of forests. However, it remains unclear whether this trend will continue, preventing for accurate projections of ecosystem functioning and climate feedbacks. Several ecophysiological mechanisms have been proposed to regulate the timing of leaf emergence in response to changing environmental cues, but the relative importance of those mechanisms remains unclear. Here, we use 727,401 direct phenological observations of common European forest trees to examine the dominant controls on leaf‐out. Using the emerging mechanisms, we forecast future trajectories of spring arrival and evaluate the consequences for forest carbon dynamics. By representing hypothesized relationships with autumn temperature, winter chilling, and the timing of spring onset, we accurately predicted reductions in the advance of leaf‐out. There was a strong consensus between our empirical model and existing process‐based models, revealing that the advance in leaf‐out will not exceed 2 weeks over the rest of the century. We further estimate that, under a ‘business‐as‐usual’ climate scenario, earlier spring arrival will enhance NPP of temperate and boreal forests by ~0.2 Gt per year at the end of the century. In contrast, previous estimates based on a simple degree‐day model range around 0.8 Gt. As such, the expected NPP is drastically reduced in our updated model relative to previous estimates—by a total of ~25 Gt over the rest of the century. These findings reveal important environmental constraints on the productivity of broad‐leaved deciduous trees and highlight that shifting spring phenology is unlikely to slow the rate of warming by offsetting anthropogenic carbon emissions.     

Northwestward range expansion of the bumblebee Bombus haematurus into Central Europe is associated with warmer winters and niche conservatism

Species range expansions are crucial for understanding niche formation and the interaction with the environment. Here, we studied the bumblebee Bombus haematurus Kriechbaumer, 1870, a species historically distributed from northern Serbia through northern Iran which has very recently started expanding northwestward into Central Europe without human-mediated dispersal (i.e., it is a natural spread). After updating the global distribution of this species, we investigated if niche shifts took place during this range expansion between newly colonized and historical areas. In addition, we have explored which climatic factors may have favored the natural range expansion of the species. Our results indicated that Bombus haematurus has colonized large territories in 7 European countries outside the historical area in the period from the 1980s to 2018, a natural expansion over an area that equals 20% of the historical distribution. In addition, this bumblebee performs generalism in flower visitation and it occurs in different habitats, although a preference for forested areas clearly emerges. The land-use associated with the species in the colonized areas is similar to the historical distribution, indicating that no major niche shifts occurred during the spread. Furthermore, in recently colonized localities, the range expansion was associated with warming temperatures during the winter and also during both queen overwintering and emergence phases. These findings document a case of natural range expansion due to environmental change rather than due to niche shifts, and specifically they suggest that warmer winters could be linked to the process of natural colonization of new areas.     

Combined effects of climate and biotic interactions on the elevational range of a phytophagous insect

1. The ranges of many species have expanded in cool regions but contracted at warm margins in response to recent climate warming, but the mechanisms behind such changes remain unclear. Particular debate concerns the roles of direct climatic limitation vs. the effects of interacting species in explaining the location of low latitude or low elevation range margins. 2. The mountains of the Sierra de Guadarrama (central Spain) include both cool and warm range margins for the black-veined white butterfly, Aporia crataegi, which has disappeared from low elevations since the 1970s without colonizing the highest elevations. 3. We found that the current upper elevation limit to A. crataegi's distribution coincided closely with that of its host plants, but that the species was absent from elevations below 900 m, even where host plants were present. The density of A. crataegi per host plant increased with elevation, but overall abundance of the species declined at high elevations where host plants were rare. 4. The flight period of A. crataegi was later at higher elevations, meaning that butterflies in higher populations flew at hotter times of year; nevertheless, daytime temperatures for the month of peak flight decreased by 6.2 degrees C per 1 km increase in elevation. 5. At higher elevations A. crataegi eggs were laid on the south side of host plants (expected to correspond to hotter microclimates), whereas at lower sites the (cooler) north side of plants was selected. Field transplant experiments showed that egg survival increased with elevation. 6. Climatic limitation is the most likely explanation for the low elevation range margin of A. crataegi, whereas the absence of host plants from high elevations sets the upper limit. This contrasts with the frequent assumption that biotic interactions typically determine warm range margins, and thermal limitation cool margins. 7. Studies that have modelled distribution changes in response to climate change may have underestimated declines for many specialist species, because range contractions will be exacerbated by mismatch between the future distribution of suitable climate space and the availability of resources such as host plants.     

Montane forest ecotones moved downslope in northeastern USA in spite of warming between 1984 and 2011

Ecotones are transition zones that form, in forests, where distinct forest types meet across a climatic gradient. In mountains, ecotones are compressed and act as potential harbingers of species shifts that accompany climate change. As the climate warms in New England, USA, high‐elevation boreal forests are expected to recede upslope, with northern hardwood species moving up behind. Yet recent empirical studies present conflicting findings on this dynamic, reporting both rapid upward ecotonal shifts and concurrent increases in boreal species within the region. These discrepancies may result from the limited spatial extent of observations. We developed a method to model and map the montane forest ecotone using Landsat imagery to observe change at scales not possible for plot‐based studies, covering mountain peaks over 39 000 km². Our results show that ecotones shifted downward or stayed stable on most mountains between 1991 and 2010, but also shifted upward in some cases (13–15% slopes). On average, upper ecotone boundaries moved down ?1.5 m yr^?1 in the Green Mountains, VT, and ?1.3 m yr^?1 in the White Mountains, NH. These changes agree with remeasured forest inventory data from Hubbard Brook Experimental Forest, NH, and suggest that processes of boreal forest recovery from prior red spruce decline, or human land use and disturbance, may swamp out any signal of climate‐mediated migration in this ecosystem. This approach represents a powerful framework for evaluating similar ecotonal dynamics in other mountainous regions of the globe.     

Spatial and temporal variations in boreal forest fire frequency in northern Alberta

Abstract. Spatial and temporal variations in fire frequency in the boreal forest of Wood Buffalo National Park (WBNP) were assessed using forest stand age, fire scar and historical data. I test the hypotheses that (1) fire frequency is higher in jack pine forests and aspen forests than in black spruce forests and white spruce forests, (2) these variations in fire frequency can be related to the mean waterbreak distance (MWD) around a site and (3) fire frequency has changed over the past 300 years. The fire cycles (the time required to burn an area equal in size to the entire study area) in jack pine forests (39 years) and in aspen forests (39 years) were significantly shorter than those in black spruce forests (78 years) and in white spruce forests (96 years). The length of the fire cycle varies inversely with the MWD around a site, and the MWD was significantly higher in jack pine and aspen forests than in black or white spruce forests. It is suggested that covariations between soil type and the MWD influence, respectively, variations in forest dominant and fire frequency. A change in fire frequency at 1860 was apparent in the fire history for all of WBNP, the black spruce dominated stands, and the near and medium MWD classes. The fire cycle estimates for these classes were all significantly shorter during the period 1750 to 1859 (fire cycles = 25–49 years) than they were in the period 1860 to 1989 (fire cycles = 59–89 years). The possible roles of changes in climate and aboriginal burning practices in causing the temporal change in fire frequency are discussed.     

Molecular and ecological plant defense responses along an elevational gradient in a boreal ecosystem

Plants have the capacity to alter their phenotype in response to environmental factors, such as herbivory, a phenomenon called phenotypic plasticity. However, little is known on how plant responses to herbivory are modulated by environmental variation along ecological gradients. To investigate this question, we used bilberry (Vaccinium myrtillus L.) plants and an experimental treatment to induce plant defenses (i.e., application of methyl jasmonate; MeJA), to observe ecological responses and gene expression changes along an elevational gradient in a boreal system in western Norway. The gradient included optimal growing conditions for bilberry in this region (ca. 500 m a.s.l.), and the plant's range limits at high (ca. 900 m a.s.l.) and low (100 m a.s.l.) elevations. Across all altitudinal sites, MeJA‐treated plants allocated more resources to herbivory resistance while reducing growth and reproduction than control plants, but this response was more pronounced at the lowest elevation. High‐elevation plants growing under less herbivory pressure but more resource‐limiting conditions exhibited consistently high expression levels of defense genes in both MeJA‐treated and untreated plants at all times, suggesting a constant state of “alert.” These results suggest that plant defense responses at both the molecular and ecological levels are modulated by the combination of climate and herbivory pressure, such that plants under different environmental conditions differentially direct the resources available to specific antiherbivore strategies. Our findings are important for understanding the complex impact of future climate changes on plant–herbivore interactions, as this is a major driver of ecosystem functioning and biodiversity.     

Wood anatomical traits in black spruce reveal latent water constraints on the boreal forest

The effects of climate change on high‐latitude forest ecosystems are complex, making forecasts of future scenarios uncertain. The predicted lengthening of the growing season under warming conditions is expected to increase tree growth rates. However, there is evidence of an increasing sensitivity of the boreal forest to drought stress. To assess the influence of temperature and precipitation on the growth of black spruce (Picea mariana), we investigated long‐term series of wood anatomical traits on 20 trees from four sites along 600 km, the latitudinal range of the closed boreal forest in Quebec, Canada. We correlated the anatomical traits resolved at intraring level with daily temperature, vapor pressure deficit (VPD), and precipitation during the 1943–2010 period. Tree‐ring width, number of cells per ring and cell wall thickness were positively affected by spring and summer daily mean and maximum temperature at the northern sites. These results agree with the well‐known positive effect of high temperatures on tree ring formation at high latitudes. However, we captured, for the first time in this region, the latent impact of water availability on xylem traits. Indeed, in all the four sites, cell lumen area showed positive correlations with daily precipitation (mostly at low latitude), and/or negative correlations with daily mean and maximum temperature and VPD (mostly at high latitude). We inferred that drought, due to high temperatures, low precipitations, or both, negatively affects cell enlargement across the closed boreal forest, including the northernmost sites. The production of tracheids with narrower lumen, potentially more resistant to cavitation, could increase xylem hydraulic safety under a warmer and drier climate. However, this would result in lower xylem conductivity, with consequent long‐term hydraulic deterioration, growth decline, and possibly lead to tree dieback, as observed in other forest ecosystems at lower latitudes.     

Unusual forest growth decline in boreal North America covaries with the retreat of Arctic sea ice

The 20th century was a pivotal period at high northern latitudes as it marked the onset of rapid climatic warming brought on by major anthropogenic changes in global atmospheric composition. In parallel, Arctic sea ice extent has been decreasing over the period of available satellite data records. Here, we document how these changes influenced vegetation productivity in adjacent eastern boreal North America. To do this, we used normalized difference vegetation index (NDVI) data, model simulations of net primary productivity (NPP) and tree‐ring width measurements covering the last 300 years. Climatic and proxy‐climatic data sets were used to explore the relationships between vegetation productivity and Arctic sea ice concentration and extent, and temperatures. Results indicate that an unusually large number of black spruce (Picea mariana) trees entered into a period of growth decline during the late‐20th century (62% of sampled trees; n = 724 cross sections of age >70 years). This finding is coherent with evidence encoded in NDVI and simulated NPP data. Analyses of climatic and vegetation productivity relationships indicate that the influence of recent climatic changes in the studied forests has been via the enhanced moisture stress (i.e. greater water demands) and autotrophic respiration amplified by the declining sea ice concentration in Hudson Bay and Hudson Strait. The recent decline strongly contrasts with other growth reduction events that occurred during the 19th century, which were associated with cooling and high sea ice severity. The recent decline of vegetation productivity is the first one to occur under circumstances related to excess heat in a 300‐year period, and further culminates with an intensifying wildfire regime in the region. Our results concur with observations from other forest ecosystems about intensifying temperature‐driven drought stress and tree mortality with ongoing climatic changes.     

Stimulation of both photosynthesis and respiration in response to warmer and drier conditions in a boreal peatland ecosystem

Peatland ecosystems have been consistent carbon (C) sinks for millennia, but it has been predicted that exposure to warmer temperatures and drier conditions associated with climate change will shift the balance between ecosystem photosynthesis and respiration providing a positive feedback to atmospheric CO₂ concentration. Our main objective was to determine the sensitivity of ecosystem photosynthesis, respiration and net ecosystem production (NEP) measured by eddy covariance, to variation in temperature and water table depth associated with interannual shifts in weather during 2004–2009. Our study was conducted in a moderately rich treed fen, the most abundant peatland type in western Canada, in a region (northern Alberta) where peatland ecosystems are a significant landscape component. During the study, the average growing season (May–October) water depth declined approximately 38 cm, and temperature [expressed as cumulative growing degree days (GDD, March–October)] varied approximately 370 GDD. Contrary to previous predictions, both ecosystem photosynthesis and respiration showed similar increases in response to warmer and drier conditions. The ecosystem remained a strong net sink for CO₂ with an average NEP (± SD) of 189 ± 47 g C m^?2 yr^?1. The current net CO₂ uptake rates were much higher than C accumulation in peat determined from analyses of the relationship between peat age and cumulative C stock. The balance between C addition to, and total loss from, the top 0–30 cm depth (peat age range 0–70 years) of shallow peat cores averaged 43 ± 12 g C m^?2 yr^?1. The apparent long‐term average rate of net C accumulation in basal peat samples was 19–24 g C m^?2 yr^?1. The difference between current rates of net C uptake and historical rates of peat accumulation is likely a result of vegetation succession and recent increases in tree establishment and productivity.     

Differential declines in Alaskan boreal forest vitality related to climate and competition

Rapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site‐level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south‐central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect‐related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early‐successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand.     

Insect defoliation modulates influence of climate on the growth of tree species in the boreal mixed forests of eastern Canada

Increasing air temperatures and changing precipitation patterns due to climate change can affect tree growth in boreal forests. Periodic insect outbreaks affect the growth trajectory of trees, making it difficult to quantify the climate signal in growth dynamics at scales longer than a year. We studied climate‐driven growth trends and the influence of spruce budworm (Choristoneura fumiferana Clem.) outbreaks on these trends by analyzing the basal area increment (BAI) of 2058 trees of Abies balsamea (L.) Mill., Picea glauca (Moench) Voss, Thuja occidentalis L., Populus tremuloides Michx., and Betula papyrifera Marsh, which co‐occurs in the boreal mixedwood forests of western Quebec. We used a generalized additive mixed model (GAMM) to analyze species‐specific trends in BAI dynamics from 1967 to 1991. The model relied on tree size, cambial age, degree of spruce budworm defoliation, and seasonal climatic variables. Overall, we observed a decreasing growth rate of the spruce budworm host species, A. balsamea and P. glauca between 1967 and 1991, and an increasing growth rate for the non‐host, P. tremuloides, B. papyrifera, and T. occidentalis. Our results suggest that insect outbreaks may offset growth increases resulting from a warmer climate. The observation warrants the inclusion of the spruce budworm defoliation into models predicting future forest productivity.     

Interactions between range‐expanding tropical fishes and the northern Gulf of Mexico red snapper Lutjanus campechanus

Experimental investigation of the intensity of potential competitive interactions among increasingly abundant tropically‐associated grey Lutjanus griseus and lane snapper Lutjanus synagris and resident northern Gulf of Mexico (nGOM) red snapper Lutjanus campechanus was undertaken in large outdoor mesocosms. In pair‐wise interaction trials, compared with L. synagris, L. campechanus demonstrated significantly increased roving behaviour and predatory activity. While no significant difference in these activities was observed between L. campechanus and L. griseus, when all three snappers (Lutjanidae) were grouped together L. campechanus swimming activity significantly decreased in the presence of both tropically‐associated species. Overall, L. campechanus were more active and aggressive predators and appear to be competitively resistant to L. griseus and L. synagris. As lower latitude species have continued to become increasingly prevalent in nGOM habitats and regional warming continues to affect resident reef‐associated fishes, these findings contribute to the assessment of the effects of warming‐related species shifts upon nGOM fishes and document current partial resilience of L. campechanus to climate‐related expansions of tropical confamilials.     

Climate warming increases biological control agent impact on a non‐target species

Climate change may shift interactions of invasive plants, herbivorous insects and native plants, potentially affecting biological control efficacy and non‐target effects on native species. Here, we show how climate warming affects impacts of a multivoltine introduced biocontrol beetle on the non‐target native plant Alternanthera sessilis in China. In field surveys across a latitudinal gradient covering their full distributions, we found beetle damage on A. sessilis increased with rising temperature and plant life history changed from perennial to annual. Experiments showed that elevated temperature changed plant life history and increased insect overwintering, damage and impacts on seedling recruitment. These results suggest that warming can shift phenologies, increase non‐target effect magnitude and increase non‐target effect occurrence by beetle range expansion to additional areas where A. sessilis occurs. This study highlights the importance of understanding how climate change affects species interactions for future biological control of invasive species and conservation of native species.