Climate controls the distribution of a widespread invasive species: implications for future range expansion

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Land use changes could modify future negative effects of climate change on old‐growth forest indicator species

Aim

Climate change is expected to have major impacts on terrestrial biodiversity at all ecosystem levels, including reductions in species‐level distribution and abundance. We aim to test the extent to which land use management, such as setting‐aside forest from production, could reduce climate‐induced biodiversity impacts for specialist species over large geographical gradients.

Location

Sweden.

Methods

We applied ensembles of species distribution models based on citizen science data for six species of red‐listed old‐forest indicator fungi confined to spruce dead wood. We tested the effect on species habitat suitabilities of alternative climate change scenarios and varying amounts of forest set‐aside from production over the coming century.

Results

With 3.6% of forest area set‐aside from production and assuming no climate change, overall habitat suitabilities for all six species were projected to increase in response to maturing spruce in set‐aside forest. However, overall habitat suitabilities for all six species were projected to decline under climate change scenario RCP4.5 (intermediate–low emissions), with even greater declines projected under RCP 8.5 (high emissions). Increasing the amount of forest set‐aside to 16% resulted in significant increases in overall habitat suitability, with one species showing an increase. A further increase to 32% forest set‐aside resulted in considerably more positive trends, with three of six species increasing.

Main conclusions

There is interspecific variation in the importance of future macroclimate and resource availability on species occurrence. However, large‐scale conservation measures, such as increasing resource availability through setting aside forest from production, could reduce future negative effects from climate change, and early investment in conservation is likely to reduce the future negative impacts of climate change on specialist species.     

Suitable areas for the invasion expansion of Xylocopa bees in South America

The introduction of exotic species into native ecosystems can be a cause for concern when those species are invasive. Invasive species cause ecological problems and have socio-cultural impacts on human health and the economy; for example, invasive bees may negatively impact their introduced ecosystem by spreading diseases or outcompeting native pollinators. Xylocopa spp. bees are diverse and distributed throughout the Neotropics. However, Xylocopa augusti (Lepeletier, 1841) and Xylocopa splendidula (Lepeletier, 1841) are not native to Mediterranean Chile. This study aimed to evaluate the invasive potential of these exotic species and predict the potential macroecological effects of their invasions. We also aimed to pinpoint possible distributions for these species throughout South America. We correlated biogeographic occurrence data with climatic variables for each species to model their potential distribution in both current and future scenarios. The models provide strong evidence that both species are changing their distributions: their ranges are expanding towards western South America, particularly Bolivia, Chile and Peru. We demonstrate an increase in niche overlap between these species and show there are new geographic areas vulnerable to the establishment of these invasive bees under current and future climate conditions. These data suggest that these bees may adapt their geographic distribution as the climate changes and pose a threat to native pollinators in new geographic areas.     

Niche conservatism and the potential for the crayfish Procambarus clarkii to invade South America

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Increasing potential risk of a global aquatic invader in Europe in contrast to other continents under future climate change

Background

Anthropogenically-induced climate change can alter the current climatic habitat of non-native species and can have complex effects on potentially invasive species. Predictions of the potential distributions of invasive species under climate change will provide critical information for future conservation and management strategies. Aquatic ecosystems are particularly vulnerable to invasive species and climate change, but the effect of climate change on invasive species distributions has been rather neglected, especially for notorious global invaders.

Methodology/Principal Findings

We used ecological niche models (ENMs) to assess the risks and opportunities that climate change presents for the red swamp crayfish (Procambarus clarkii), which is a worldwide aquatic invasive species. Linking the factors of climate, topography, habitat and human influence, we developed predictive models incorporating both native and non-native distribution data of the crayfish to identify present areas of potential distribution and project the effects of future climate change based on a consensus-forecast approach combining the CCCMA and HADCM3 climate models under two emission scenarios (A2a and B2a) by 2050. The minimum temperature from the coldest month, the human footprint and precipitation of the driest quarter contributed most to the species distribution models. Under both the A2a and B2a scenarios, P. clarkii shifted to higher latitudes in continents of both the northern and southern hemispheres. However, the effect of climate change varied considerately among continents with an expanding potential in Europe and contracting changes in others.

Conclusions/Significance

Our findings are the first to predict the impact of climate change on the future distribution of a globally invasive aquatic species. We confirmed the complexities of the likely effects of climate change on the potential distribution of globally invasive species, and it is extremely important to develop wide-ranging and effective control measures according to predicted geographical shifts and changes.     

Forest pattern,climate and vulcanism in central North Island,New Zealand

Abstract. A quantitative study of relationships between forest pattern and environment in the central North Island, New Zealand, is based on forest composition data from ca. 2000 existing plots distributed throughout the forests of the region. Estimates of mean annual temperature, rainfall, and solar radiation are derived for each plot from mathematical surfaces fitted to climate station data. Estimates of the depth of the last major rhyolitic eruption, (Taupo Pumice, ca. 130 AD) are derived from isopach maps. A classification procedure is used to identify broad compositional groups. Generalised linear models are used to examine relationships between major species and climatic and other physical factors. Significant relationships are identified between the distributions of both plot groups and species, and climate, vulcanism, topography and drainage. Among these factors, temperature and/or solar radiation are indicated as major determinants of the regional forest pattern, with rainfall, topography, and drainage acting at a secondary level. The role of the Taupo Pumice eruption is more difficult to interpret, and its effects seem to have been greatly influenced by topography. Deep extensive deposits of tephra on flat-to-rolling sites close to the eruption centre have probably favoured the current dominance of these sites by more rapidly dispersing conifers. In contrast, on adjacent steep sites where forest destruction was likely to be less severe, slow-dispersing Nothofagus species are largely dominant. Further work is needed to understand the factors favouring conifer dominance of the central basins and the degree to which Nothofagus species might expand their range in the future.     

Selection for commercial forestry determines global patterns of alien conifer invasions

Question Are the patterns of alien conifer (Pinaceae, Cupressaceae) invasions different between continents, and how is invasion success influenced by commercial forestry practices? Location Temperate and subtropical countries and regions (n = 60) from five continents spanning both hemispheres. Methods We used generalized linear mixed models to test how continent identity, region area and use in commercial forestry affect probabilities of Pinaceae and Cupressaceae species to escape following introduction and cumulative logit regression models to assess how these predictors affect the likelihood that a species becomes naturalized or invasive. Results Sixty Pinaceae of a global total of 232 and 26 Cupressaceae of a total of 142 species have escaped from cultivation across the study regions examined. Average numbers of both alien Pinaceae and Cupressaceae species per region were highest in Oceania, followed by Africa. Moreover, the probability of alien Cupressaceae and Pinaceae becoming naturalized or invasive was particularly high in these two continents. For both families, species used in commercial forestry have a significantly higher probability of escape than those which are only introduced for ornamental or other purposes. In the case of Pinaceae, forestry species also become naturalized or invasive more frequently than non‐forestry species, while no such effect was detectable for Cupressaceae. Conclusions We found that non‐native conifers are more likely to escape from cultivation, naturalize and turn into invasive weeds on the continents of the Southern Hemisphere. In addition to this biogeographic signal, introduction effort strongly determines the behaviour of introduced Pinaceae, and less so, Cupressaceae. A clear conflict exists between the economic benefits of conifer forestry and the risks to the environment from invasions. Future expansion of commercial forestry should address spatial planning to ecosystems vulnerable to invasion and adopt comprehensive risk assessment procedures.     

Native vegetation structure,landscape features and climate shape non-native plant richness and cover in New Zealand native shrublands

Aim

Studies investigating the determinants of plant invasions rarely examine multiple factors and often only focus on the role played by native plant species richness. By contrast, we explored how vegetation structure, landscape features and climate shape non-native plant invasions across New Zealand in mānuka and kānuka shrublands.

Location

New Zealand.

Method

We based our analysis on 247 permanent 20 × 20-m plots distributed across New Zealand surveyed between 2009 and 2014. We calculated native plant species richness and cumulative cover at ground, understorey and canopy tiers. We examined non-native species richness and mean species ground cover in relation to vegetation structure (native richness and cumulative cover), landscape features (proportion of adjacent anthropogenic land cover, distance to nearest road or river) and climate. We used generalized additive models (GAM) to assess which variables had greatest importance in determining non-native richness and mean ground cover and whether these variables had a similar effect on native species in the ground tier.

Results

A positive relationship between native and non-native plant species richness was not due to their similar responses to the variables examined in this study. Higher native canopy richness resulted in lower non-native richness and mean ground cover, whereas higher native ground richness was associated with higher native canopy richness. Non-native richness and mean ground cover increased with the proportion of adjacent anthropogenic land cover, whereas for native richness and mean ground cover, this relationship was negative. Non-native richness increased in drier areas, while native richness was more influenced by temperature.

Main Conclusions

Adjacent anthropogenic land cover seems to not only facilitate non-native species arrival by being a source of propagules but also aids their establishment as a result of fragmentation. Our results highlight the importance of examining both cover and richness in different vegetation tiers to better understand non-native plant invasions.     

Can Invasive Species Facilitate Native Species? Evidence of How, When, and Why These Impacts Occur

Although the predatory and competitive impacts of biological invasions are well documented, facilitation of native species by non-indigenous species is frequently overlooked. A search through recent ecological literature found that facilitative interactions between invasive and native species occur in a wide range of habitats, can have cascading effects across trophic levels, can re-structure communities, and can lead to evolutionary changes. These are critical findings that, until now, have been mostly absent from analyses of ecological impacts of biological invasions. Here I present evidence for several mechanisms that exemplify how exotic species can facilitate native species. These mechanisms include habitat modification, trophic subsidy, pollination, competitive release, and predatory release. Habitat modification is the most frequently documented mechanism, reflecting the drastic changes generated by the invasion of functionally novel habitat engineers. Further, I predict that facilitative impacts on native species will be most likely when invasive species provide a limiting resource, increase habitat complexity, functionally replace a native species, or ameliorate predation or competition. Finally, three types of facilitation (novel, substitutive, and indirect) define why exotic species can lead to facilitation of native species. It is evident that understanding biological invasions at the community and ecosystem levels will be more accurate if we integrate facilitative interactions into future ecological research. Since facilitative impacts of biological invasions can occur with native endangered species, and can have wide-ranging impacts, these results also have important implications for management, eradication, and restoration.Contribution Number 2293, Bodega Marine Laboratory, University of California at Davis.     

Oeceoclades maculata, an alien tropical orchid in a Caribbean rainforest

Background and Aims

Undisturbed forest habitat can be relatively impenetrable to invasive, non-native species. Orchids are not commonly regarded as invasive, but some species have become invasive and these generally depend on habitat disturbance. One of the most aggressive orchids is Oeceoclades maculata, a terrestrial species with remarkable ecological amplitude. Originally from tropical Africa, it is now widespread in the neotropics. By associating its local distribution with land-use history and habitat characteristics, it was determined whether O. maculata is dependent on habitat disturbance. It was also investigated whether this exotic orchid occupies the same habitat space as two sympatric native species.

Methods

Six 10 m × 500 m transects were censused in June 2007 on the 16-ha Luquillo Forest Dynamics Plot, located in the Luquillo Mountains, Puerto Rico. The plot had been mapped for historical land use, topography and soil type.

Key Results

Oeceoclades maculata was the most abundant of three orchid species surveyed and was found in all four historical cover classes. In cover class 3 (50–80 % forest cover in 1936), 192 of 343 plants were found at a density of 0·48 plants per 5 × 5 m subplot. Over 93 % of the 1200 subplots surveyed were composed of Zarzal or Cristal soil types, and O. maculata was nearly evenly distributed in both. The orchid was most common on relatively flat terrain. The distribution and abundance of two sympatric orchid species were negatively associated with that of the invasive species.

Conclusions

Oeceoclades maculata does penetrate ‘old growth’ forest but is most abundant in areas with moderate levels of past disturbance. Soil type makes little difference, but slope of terrain can be important. The negative association between O. maculata and native species may reflect differences in habitat requirements or a negative interaction perhaps at the mycorrhizal level.Key words: Oeceoclades maculata, Wullschlaegelia calcarata, Prescottia stachyodes, Orchidaceae, land-use history, tropical forest disturbance, terrestrial orchids, invasive species, Luquillo Experimental Forest, Puerto Rico, forest recovery, Caribbean     

Multiple stressors facilitate the spread of a non‐indigenous bivalve in the Mediterranean Sea

Aim

The introduction of non‐indigenous species (NIS) via man‐made corridors connecting previously disparate oceanic regions is increasing globally. However, the environmental and anthropogenic factors facilitating invasion dynamics and their interactions are still largely unknown. This study compiles and inputs available data for the NIS bivalve Brachidontes pharaonis across the invaded biogeographic range in the Mediterranean basin into a species distribution model to predict future spread under a range of marine scenarios.

Location

Mediterranean Sea.

Methods

A systematic review produced the largest presence database ever assembled to inform the selection of biological, chemical and physical factors linked to the spread of B. pharaonis through the Suez Canal. We carried out a sensitivity analysis to simulate current and future trophic and salinity scenarios. A species distribution model was run to determine key drivers of invasion, quantify interactive impacts arising from a range of trophic states, salinity conditions and climatic scenarios and forecast future trajectories for the spread of NIS into new regions under multiple‐parameter scenarios (based on the main factors identified from the systematic review).

Results

Impacts on invasion trajectory arising from climate change and interactions with increasing salinity from the new opening of the canal were the primary drivers of expansion across the basin, the effects of which were further enhanced by eutrophication. Predictions of the current distribution were most accurate when multiple stressors were used to drive the model. A habitat suitability index developed at a subcontinental scale from model outputs identified novel favourable conditions for future colonization at specific locations under 2030 and 2050 climatic scenarios.

Main conclusions

Future expansion of B. pharaonis will be enhanced by climate‐facilitated increased sea temperature, interacting with increasing pressures from salinity and eutrophication. The spatially explicit risk output maps of invasions represent a powerful visual product for use in communication of the spread of NIS and decision‐support tools for scientists and policymakers. The suggested approach, the observed distribution pattern and driving processes can be applied to other NIS species and regions by providing novel forecasts of species occurrences under future multiple stressor scenarios and the location of suitable recipient habitats with respect to anthropogenic and environmental parameters.     

Species Invasiveness in Biological Invasions: A Modelling Approach

The study of invasiveness, the traits that enable a species to invade a habitat, and invasibility, the habitat characteristics that determine its susceptibility to the establishment and spread of an invasive species, provide a useful conceptual framework to formulate the biological invasion problem in a modelling context. Another important aspect is the complex interaction emerging among the invader species, the noninvader species already present in the habitat, and the habitat itself. Following a modelling approach to the biological invasion problem, we present a spatially explicit cellular automaton model (Interacting Multiple Cellular Automata (IMCA)). We use field parameters from the invader Gleditsia triacanthos and the native Lithraea ternifolia in montane forests of central Argentina as a case study to compare outputs and performance of different models. We use field parameters from another invader, Ligustrum lucidum, and the native Fagara coco from the same system to run the cellular automaton model. We compare model predictions with invasion values from aerial photographs. We discuss in detail the importance of factors affecting species invasiveness, and give some insights into habitat invasibility and the role of interactions between them. Finally, we discuss the relevance of mathematical modelling for studying and predicting biological invasions. The IMCA model provided a suitable context for integrating invasiveness, invasibility, and the interactions. In the invasion system studied, the presence of an invader's juvenile bank not only accelerated the rate of invasion but was essential to ensure invasion. Using the IMCA model, we were able to determine that not only adult survival but particularly longevity of the native species influenced the spread velocity of the invader, at least when a juvenile bank is present. Other factors determining velocity of invasion detected by the IMCA model were seed dispersal distance and age of reproductive maturity. We derived relationships between species' adult survival, fecundity and longevity of both theoretical and applied relevance for biological invasions. Invasion velocities calculated from the aerial photographs agreed well with predictions of the IMCA model.     

Different climatic envelopes among invasive populations may lead to underestimations of current and future biological invasions

Aim We explore the impact of calibrating ecological niche models (ENMs) using (1) native range (NR) data versus (2) entire range (ER) data (native and invasive) on projections of current and future distributions of three Hieracium species. Location H. aurantiacum, H. murorum and H. pilosella are native to Europe and invasive in Australia, New Zealand and North America. Methods Differences among the native and invasive realized climatic niches of each species were quantified. Eight ENMs in BIOMOD were calibrated with (1) NR and (2) ER data. Current European, North American and Australian distributions were projected. Future Australian distributions were modelled using four climate change scenarios for 2030. Results The invasive climatic niche of H. murorum is primarily a subset of that expressed in its native range. Invasive populations of H. aurantiacum and H. pilosella occupy different climatic niches to those realized in their native ranges. Furthermore, geographically separate invasive populations of these two species have distinct climatic niches. ENMs calibrated on the realized niche of native regions projected smaller distributions than models incorporating data from species’ entire ranges, and failed to correctly predict many known invasive populations. Under future climate scenarios, projected distributions decreased by similar percentages, regardless of the data used to calibrate ENMs; however, the overall sizes of projected distributions varied substantially. Main conclusions This study provides quantitative evidence that invasive populations of Hieracium species can occur in areas with different climatic conditions than experienced in their native ranges. For these, and similar species, calibration of ENMs based on NR data only will misrepresent their potential invasive distribution. These errors will propagate when estimating climate change impacts. Thus, incorporating data from species’ entire distributions may result in a more thorough assessment of current and future ranges, and provides a closer approximation of the elusive fundamental niche.     

Do invasive freshwater fish species grow better when they are invasive?

A frequent assumption in invasion ecology is that invasive species have enhanced growth rates in their invasive ranges. However, invasions frequently occur in sub‐tropical and tropical environments where growth could be higher simply due to climatic conditions rather than novel habitat. In this study, a meta‐analysis of growth rates (length‐at‐age data) was completed for six invasive freshwater fish species: common carp Cyprinus carpio, largemouth bass Micropterus salmoides, brown trout Salmo trutta, brown bullhead Ictalurus nebulosus, flathead catfish Pylodictis olivaris and northern snakehead Channa argus. Significant effects of climate on growth were observed for all species except common carp, and following normalization of growth for climate effects, a range of growth responses between native and invasive populations were revealed. Two species (brown trout, flathead catfish) showed significantly increased growth rates in invasive compared to native ranges, but two species (common carp, largemouth bass) showed significantly faster growth in native ranges, and two other species (northern snakehead, brown bullhead) showed no difference in growth rates. No species showed both significantly enhanced growth rates and initial sizes in invasive compared to native ranges. Using the comparative method, countergradient growth variations were apparent for all species within their native ranges and for all but one species in invasive ranges. Invasive populations of freshwater fish do not always grow faster when invasive and future studies need to consider growth covariates (e.g. climate and countergradient growth) prior to comparing life‐history differences between invasive and native populations.     

The abundance,distribution and structural characteristics of tree‐holes in Nothofagus forest,New Zealand

Tree‐holes provide an important microhabitat that is used for feeding, roosting and breeding by numerous species around the world. Yet despite their ecological importance for many of New Zealand's endangered species, few studies have investigated the abundance or distribution of tree‐holes in native forests. We used complementary ground and climbed tree surveys to determine the abundance, distribution and characteristics of tree‐holes in undisturbed Nothofagus forest in the Lewis Pass, New Zealand. We found that hole‐bearing trees were surprisingly abundant compared with many other studies, including Australian Eucalyptus species and American beech. In fact, we estimated as many as 3906 tree‐holes per hectare, of which 963 holes per hectare were potentially large enough to provide roost sites for hole‐nesting bats in New Zealand, while only eight holes per hectare were potentially suitable for specialist hole‐nesting birds. This was of great interest as primary cavity‐excavating animals are absent from New Zealand forests, compared with North America and Australia. Moreover, tree‐hole formation in New Zealand is likely to be dominated by abiotic processes, such as branch breakage from windstorms and snow damage. As has been found in many other studies, tree‐holes were not uniformly distributed throughout the forest. Tree‐holes were significantly more abundant on the least abundant tree species, Nothofagus fusca, than on either N. menziesii or N. solandri. In addition to tree species, tree size was also an important factor influencing the structural characteristics of tree‐holes and their abundance in this forest. Moreover, these trends were not fully evident without climbed tree surveys. Our results revealed that ground‐based surveys consistently underestimated the number of tree‐holes present on Nothofagus trees, and illustrate the importance of using climbed inspections where possible in tree‐hole surveys. We compare our results with other studies overseas and discuss how these are linked to the biotic and abiotic processes involved in tree‐hole formation. We consider the potential implications of our findings for New Zealand's hole‐dwelling fauna and how stand dynamics and past and future forest management practices will influence the structural characteristics of tree‐holes and their abundance in remnant forest throughout New Zealand.     

The potential global distribution of the Bronze bug Thaumastocoris peregrinus Carpintero and Dellapé (Hemiptera: Thaumastocoridae)

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Invasive fountain grass (Pennisetum setaceum (Forssk.) Chiov.) increases its potential area of distribution in Tenerife island under future climatic scenarios

Mapping the distribution of invasive species under current and future climate conditions is crucial to implement sustainable and effective conservation strategies. Several studies showed how invasive species may benefit from climate change fostering their invasion rate and, consequently, affecting the native species community. In the Canary Islands and on Tenerife in particular, previous research mostly focused on climate change impacts on the native communities, whereas less attention has been paid on alien species distribution under climate change scenarios. In this study, we modelled the habitat distribution of Pennisetum setaceum, one of the most invasive alien species on Tenerife. In addition, we described the species’ potential distribution shift in the light of two climate change scenarios (RCP2.6, RCP8.5), highlighting the areas that should be prioritized during management and eradication programs. P. setaceum’s suitable areas are located in the coastal area, with higher habitat suitability near cities and below 800 m asl. In both future climate change scenarios, the geographic distribution of P. setaceum suitable areas is characterized by an elevational shift, which is more pronounced in the RCP8.5 scenario. Despite being drought resistant, water supply is crucial for the species’ seed germination, thus supporting future species’ shift to higher elevation and in the north–north–west part of the island, where it could benefit from the combined effect of orographic precipitations and humidity carried by trade winds.

     

Severity of impacts of an introduced species corresponds with regional eco‐evolutionary experience

Invasive plant impacts vary widely across introduced ranges. We tested the hypothesis that differences in the eco‐evolutionary experience of native communities with the invader correspond with the impacts of invasive species on native vegetation, with impacts increasing with ecological novelty. We compared plant species richness and composition beneath Pinus contorta to that in adjacent vegetation and other P. contorta stands across a network of sites in its native (Canada and USA) and non‐native (Argentina, Chile, Finland, New Zealand, Scotland, Sweden) ranges. At sites in North America and Europe, within the natural distribution of the genus Pinus, P. contorta was not associated with decreases in diversity. In the Southern Hemisphere, where there are no native Pinaceae, plant communities beneath P. contorta were less diverse than in other regions and compared to uninvaded native vegetation. Effects on native vegetation were particularly pronounced where P. contorta was a more novel life form and exhibited higher growth rates. Our results support the hypothesis that the eco‐evolutionary experience of the native vegetation, and thus the novelty of the invader, determines the magnitude of invader impacts on native communities. Understanding the eco‐evolutionary context of invasions will help to better understand and predict where invasion impacts will be greatest and to prioritize invasive species management.     

Influence of bioclimatic variables on tree-line conifer distribution in the Greater Yellowstone Ecosystem: implications for species of conservation concern

Aim Tree‐line conifers are believed to be limited by temperature worldwide, and thus may serve as important indicators of climate change. The purpose of this study was to examine the potential shifts in spatial distribution of three tree‐line conifer species in the Greater Yellowstone Ecosystem under three future climate‐change scenarios and to assess their potential sensitivity to changes in both temperature and precipitation. Location This study was performed using data from 275 sites within the boundaries of Yellowstone and Grand Teton national parks, primarily located in Wyoming, USA. Methods We used data on tree‐line conifer presence from the US Forest Service Forest Inventory and Analysis Program. Climatic and edaphic variables were derived from spatially interpolated maps and approximated for each of the sites. We used the random‐forest prediction method to build a model of predicted current and future distributions of each of the species under various climate‐change scenarios. Results We had good success in predicting the distribution of tree‐line conifer species currently and under future climate scenarios. Temperature and temperature‐related variables appeared to be most influential in the distribution of whitebark pine (Pinus albicaulis), whereas precipitation and soil variables dominated the models for subalpine fir (Abies lasiocarpa) and Engelmann spruce (Picea engelmannii). The model for whitebark pine substantially overpredicted absences (as compared with the other models), which is probably a result of the importance of biological factors in the distribution of this species. Main conclusions These models demonstrate the complex response of conifer distributions to changing climate scenarios. Whitebark pine is considered a ‘keystone’ species in the subalpine forests of western North America; however, it is believed to be nearly extinct throughout a substantial portion of its range owing to the combined effects of an introduced pathogen, outbreaks of the native mountain pine beetle (Dendroctonus ponderosae), and changing fire regimes. Given predicted changes in climate, it is reasonable to predict an overall decrease in pine‐dominated subalpine forests in the Greater Yellowstone Ecosystem. In order to manage these forests effectively with respect to future climate, it may be important to focus attention on monitoring dry mid‐ and high‐elevation forests as harbingers of long‐term change.