Elevational trends in butterfly phenology: implications for species responses to climate change

1. Impacts of global change on the distribution, abundance, and phenology of species have been widely documented. In particular, recent climate change has led to widespread changes in animal and plant seasonality, leading to debate about its potential to cause phenological mismatches among interacting taxa. 2. In mountainous regions, populations of many species show pronounced phenological gradients over short geographic distances, presenting the opportunity to test for effects of climate on phenology, independent of variation in confounding factors such as photoperiod. 3. Here we show for 32 butterfly species sampled for five years over a 1700 m gradient (560–2260 m) in a Mediterranean mountain range that, on average, annual flight period is delayed with elevation by 15–22 days per kilometre. Species mainly occurring at low elevations in the region, and to some extent those flying earlier in the year, showed phenological delays of 23–36 days per kilometre, whereas the flight periods of species that occupy high elevations, or fly in late summer, were consistently more synchronised over the elevation gradient. 4. Elevational patterns in phenology appear to reflect a narrowing phenological window of opportunity for larval and adult butterfly activity of high elevation and late‐flying species. 5. Here, we speculate as to the causes of these patterns, and the consequences for our ability to predict species responses to climate change. Our results raise questions about the use of space–time substitutions in predicting phenological responses to climate change, since traits relating to flight period and environmental associations may influence the capacity of species to adapt to changing climates.     

Forest composition in Mediterranean mountains is projected to shift along the entire elevational gradient under climate change

Aim Species distribution models have been used frequently to assess the effects of climate change on mountain biodiversity. However, the value and accuracy of these assessments have been hampered by the use of low‐resolution data for species distributions and climatic conditions. Herein we assess potential changes in the distribution and community composition of tree species in two mountainous regions of Spain under specific scenarios of climate change using data with a high spatial resolution. We also describe potential changes in species distributions and tree communities along the entire elevational gradient. Location Two mountain ranges in southern Europe: the Central Mountain Range (central west of the Iberian Peninsula), and the Iberian Mountain Range (central east). Methods We modelled current and future distributions of 15 tree species (Eurosiberian, sub‐Mediterranean and Mediterranean species) as functions of climate, lithology and availability of soil water using generalized linear models (logistic regression) and machine learning models (gradient boosting). Using multivariate ordination of a matrix of presence/absence of tree species obtained under two Intergovernmental Panel on Climate Change (IPCC) scenarios (A2 and B2) for two different periods in the future (2041–70 and 2071–2100), we assessed the predicted changes in the composition of tree communities. Results The models predicted an upward migration of communities of Mediterranean trees to higher elevations and an associated decline in communities of temperate or cold‐adapted trees during the 21st century. It was predicted that 80–99% of the area that shows a climate suitable for cold–wet‐optimum Eurosiberian coniferous and broad‐leaved species will be lost. The largest overall changes were predicted for Mediterranean species found currently at low elevations, such as Pinus halepensis, Pinus pinaster, Quercus ilex ssp. ballota and Juniperus oxycedrus, with sharp increases in their range of 350%. Main conclusions It is likely that areas with climatic conditions suitable for cold‐adapted species will decrease significantly under climate warming. Large changes in species ranges and forest communities might occur, not only at high elevations within Mediterranean mountains but also along the entire elevational gradient throughout this region, particularly at low and mid‐elevations. Mediterranean mountains might lose their key role as refugia for cold‐adapted species and thus an important part of their genetic heritage.     

Shifting and extension of phenological periods with increasing temperature along elevational transects in southern Bavaria

The impact of global warming on phenology has been widely studied, and almost consistently advancing spring events have been reported. Especially in alpine regions, an extraordinary rapid warming has been observed in the last decades. However, little is known about phenological phases over the whole vegetation period at high elevations. We observed 12 phenological phases of seven tree species and measured air temperature at 42 sites along four transects of about 1000 m elevational range in the years 2010 and 2011 near Garmisch‐Partenkirchen, Germany. Site‐ and species‐specific onset dates for the phenological phases were determined and related to elevation, temperature lapse rates and site‐specific temperature sums. Increasing temperatures induced advanced spring and delayed autumn phases, in which both yielded similar magnitudes. Delayed leaf senescence could therefore have been underestimated until now in extending the vegetation period. Not only the vegetation period, but also phenological periods extended with increasing temperature. Moreover, sensitivity to elevation and temperature strongly depends on the specific phenological phase. Differences between species and groups of species (deciduous, evergreen, high elevation) were found in onset dates, phenological response rates and also in the effect of chilling and forcing temperatures. Increased chilling days highly reduced forcing temperature requirements for deciduous trees, but less for evergreen trees. The problem of shifted species associations and phenological mismatches due to species‐specific responses to increasing temperature is a recent topic in ecological research. Therefore, we consider our findings from this novel, dense observation network in an alpine area of particular importance to deepen knowledge on phenological responses to climate change.     

An elevational shift in butterfly species richness and composition accompanying recent climate change

The geographic ranges of many species have shifted polewards and uphill in elevation associated with climate warming, leading to increases in species richness at high latitudes and elevations. However, few studies have addressed community‐level responses to climate change across the entire elevational gradients of mountain ranges, or at warm lower latitudes where ecological diversity is expected to decline. Here, we show uphill shifts in butterfly species richness and composition in the Sierra de Guadarrama (central Spain) between 1967–1973 and 2004–2005. Butterfly communities with comparable species compositions shifted uphill by 293 m (± SE 26), consistent with an upward shift of approximately 225 m in mean annual isotherms. Species richness had a humped relationship with elevation, but declined between surveys, particularly at low elevations. Changes to species richness and composition primarily reflect the loss from lower elevations of species whose regional distributions are restricted to the mountains. The few colonizations by specialist low‐elevation species failed to compensate for the loss of high‐elevation species, because there are few low‐elevation species in the region and the habitat requirements of some of these prevent them from colonizing the mountain range. As a result, we estimated a net decline in species richness in approximately 90% of the region, and increasing community domination by widespread species. The results suggest that climate warming, combined with habitat loss and other drivers of biological change, could lead to significant losses in ecological diversity in mountains and other regions where species encounter their lower latitudinal‐range margins.     

Who flies first? – habitat‐specific phenological shifts of butterflies and orthopterans in the light of climate change: a case study from the south‐east Mediterranean

1. Insects undergo phenological change at different rates, showing no consistent trend between habitats, time periods, species or groups. Understanding how and why this variability occurs is crucial. 2. Phenological patterns of butterflies and Orthoptera were analysed using a novel approach of standardised major axis (SMA) analysis. It was investigated whether: (i) phenology (the mean date and duration of flight) of butterflies and Orthoptera changed from one survey (1998 and 1999 respectively) to another (2011), (ii) the rate at which phenology changed differed between taxa and (iii) phenological change was significantly different across habitat types (agriculture fields, grasslands, and forests). Using the 2011 dataset, we investigated relationships between habitat‐specific variables and species phenology. 3. For both groups, late‐emerging species had an advanced onset on the second survey while the duration showed no consistent trend for butterflies and did not change for Orthoptera. Although the rate at which phenology changed was consistent between the two groups, at the habitat level, a longer duration of flight period emerged for butterflies in agriculture fields while Orthoptera showed no differentiation in flight duration between habitats. We found an earlier emergence of butterflies in grasslands compared to forests, attributed to habitat‐specific temperature, whereas spatial variation in humidity had a significantly lower effect on butterflies' phenology in grasslands compared to forests. A gradual delay of butterfly appearances as the canopy cover increased was also found. 4. The utility of SMA analysis was demonstrated in phenological studies and evidence was detected that both habitat type and habitat‐specific variables refine species' phenological responses.     

The influence of life history traits on the phenological response of British butterflies to climate variability since the late‐19th century

Many species of plants and animals have advanced their phenology in response to climate warming in recent decades. Most of the evidence available for these shifts is based on data from the last few decades, a period coinciding with rapid climate warming. Baseline data is required to put these recent phenological changes in a long‐term context. We analysed the phenological response of 51 resident British butterfly species using data from 83 500 specimens in the collections of the Natural History Museum, London, covering the period 1880–1970. Our analysis shows that only three species significantly advanced their phenology between 1880 and 1970, probably reflecting the relatively small increase in spring temperature over this period. However, the phenology of all but one of the species we analysed showed phenological sensitivity to inter‐annual climate variability and a significant advancement in phenology in years in which spring or summer temperatures were warm and dry. The phenologies of butterfly species were more sensitive to weather if the butterfly species was early flying, southerly distributed, and a generalist in terms of larval diet. This observation is consistent with the hypothesis that species with greater niche breadth may be more phenologically sensitive than species with important niche constraints. Comparison of our results with post‐1976 data from the UK Butterfly Monitoring Scheme show that species flying early in the year had a greater rate of phenological advancement prior to the mid‐1970s. Additionally, prior to the mid‐1970s, phenology was influenced by temperatures in March or April, whereas since 1976, February temperature had a stronger influence on the phenology. These results suggest that early flying species may be approaching the limits of phenological advancement in response to recent climate warming.     

Using a phenological network to assess weather influences on first appearance of butterflies in the Netherlands

Phenological responses of butterflies to temperature have been demonstrated in several European countries by using data from standardized butterfly monitoring schemes. Recently, phenological networks have enabled volunteers to record phenological observations at project websites. In this study, the quality of the first appearance data of butterflies from the Dutch phenological network ‘De Natuurkalender’ was examined and these data were then used to analyze trends in butterfly appearance between 2001 and 2013, the effects of climatic factors on appearance of butterflies as well as the phenological interaction of one butterfly species, Anthocharis cardamines, and its two major host plants. Although phenological networks are relatively unstructured, this study shows that data from De Natuurkalender were highly comparable to more standardized data collected by the Dutch Butterfly Monitoring Scheme. No trend in first appearance of any butterfly species was found during the time period 2001–2013. The first appearance dates of most butterflies showed, however, a clear relationship with spring temperature. Higher temperatures, especially in March and April, advanced the first appearance of butterflies. Therefore, with climatic warming in the future, earlier appearance of butterflies is expected. Although climate warming is a potential threat for phenological mismatches between different trophic levels, this study shows a similar temperature response of A. cardamines and its main host plants in the Netherlands. However, as only few phenological interactions between species are examined, further research including rarer monophagous butterfly species and their host plants is needed.     

Phenology of British butterflies and climate change

Data from a national butterfly monitoring scheme were analysed to test for relationships between temperature and three phenological measures, duration of flight period and timing of both first and peak appearance. First appearances of most British butterflies has advanced in the last two decades and is strongly related to earlier peak appearance and, for multibrooded species, longer flight period. Mean dates of first and peak appearance are examined in relation to Manley's central England temperatures, using regression techniques. We predict that, in the absence of confounding factors, such as interactions with other organisms and land‐use change, climate warming of the order of 1 °C could advance first and peak appearance of most butterflies by 2–10 days.     

Temperature‐related shifts in butterfly phenology depend on the habitat

Many species are becoming active earlier in the season as the climate becomes warmer. In parallel to phenological responses to climate change, many species have also been affected by habitat changes due to anthropogenic land use. As habitat type can directly affect microclimatic conditions, concurrent changes in climate and habitat could have interacting effects on the phenology of species. Temperature‐related shifts in phenology, however, have mostly been studied independent of habitat types. Here, I used long‐term data from a highly standardized monitoring program with 519 transects to study how phenology of butterflies is affected by ambient temperature and habitat type. I compared forests, agricultural areas and settlements, reflecting three major land use forms, and considered butterfly species that were observed in all three of these habitats. Seasonal appearance of the butterflies was affected both by the ambient temperature and the habitat type. As expected, warmer temperatures led to an overall advancement of the appearance and flight period of most species. Surprisingly, however, phenology of species was delayed in settlement habitats, even though this habitat type is generally associated with higher temperatures. A possible explanation is dispersal among habitat types, such that source–sink effects affect local phenology. When there is little productivity in settlement areas, observed butterflies may have immigrated from forest or agricultural habitats and thus appear later in settlements. My findings suggest that a spillover of individuals among habitats may affect phenology trends and indicate that phenological studies need to be interpreted in the context of habitat types. This becomes especially important when defining strategies to prevent or mitigate effects of climate and land‐use changes on phenology and abundance of species.     

Effects of climatic change on the phenology of butterflies in the northwest Mediterranean Basin

Phenological changes in response to climatic warming have been detected across a wide range of organisms. Butterflies stand out as one of the most popular groups of indicators of climatic change, given that, firstly, they are poikilothermic and, secondly, have been the subject of thorough monitoring programmes in several countries for a number of decades. Here we provide for the first time strong evidence of phenological change as a consequence of recent climatic warming in butterflies at a Spanish site in the northwest Mediterranean Basin. By means of the widely used Butterfly Monitoring Scheme methodology, three different phenological parameters were analysed for the most common species to test for trends over time and relationships with temperature and precipitation. Between 1988 and 2002, there was a tendency for earlier first appearance dates in all 17 butterfly species tested, and significant advances in mean flight dates in 8 out of 19 species. On the other hand, the shape of the curve of adult emergence did not show any regular pattern. These changes paralleled an increase of 1–1.5°C in mean February, March and June temperatures. Likewise, a correlation analysis indicated the strong negative effect of spring temperature on phenological parameters (i.e. higher temperatures tended to produce phenological advances), and the opposite effect of precipitation in certain months. In addition, there was some evidence to indicate that phenological responses may differ between taxonomic lineages or species with similar diets. We discuss the consequences that these changes may have on species' population abundances, especially given the expected increase in aridity in the Mediterranean Basin caused by current climatic warming. We predict that varying degrees of phenological flexibility may account for differences in species' responses and, for multivoltine species, predict strong selection favouring local seasonal adaptations such as diapause phenomena or migratory behaviour.     

Prairie plant phenology driven more by temperature than moisture in climate manipulations across a latitudinal gradient in the Pacific Northwest,USA

Plant phenology will likely shift with climate change, but how temperature and/or moisture regimes will control phenological responses is not well understood. This is particularly true in Mediterranean climate ecosystems where the warmest temperatures and greatest moisture availability are seasonally asynchronous. We examined plant phenological responses at both the population and community levels to four climate treatments (control, warming, drought, and warming plus additional precipitation) embedded within three prairies across a 520 km latitudinal Mediterranean climate gradient within the Pacific Northwest, USA. At the population level, we monitored flowering and abundances in spring 2017 of eight range‐restricted focal species planted both within and north of their current ranges. At the community level, we used normalized difference vegetation index (NDVI) measured from fall 2016 to summer 2018 to estimate peak live biomass, senescence, seasonal patterns, and growing season length. We found that warming exerted a stronger control than our moisture manipulations on phenology at both the population and community levels. Warming advanced flowering regardless of whether a species was within or beyond its current range. Importantly, many of our focal species had low abundances, particularly in the south, suggesting that establishment, in addition to phenological shifts, may be a strong constraint on their future viability. At the community level, warming advanced the date of peak biomass regardless of site or year. The date of senescence advanced regardless of year for the southern and central sites but only in 2018 for the northern site. Growing season length contracted due to warming at the southern and central sites (~3 weeks) but was unaffected at the northern site. Our results emphasize that future temperature changes may exert strong influence on the timing of a variety of plant phenological events, especially those events that occur when temperature is most limiting, even in seasonally water‐limited Mediterranean ecosystems.     

The roles of shifting and filtering in generating community‐level flowering phenology

Plant phenologies are key components of community assembly and ecosystem function, yet we know little about how phenological patterns differ among ecosystems. Community‐level phenological patterns may be driven by the filtering of species into communities based on their phenology or by intraspecific responses to local conditions that shift when species flower. To understand the relative roles of filtering and shifting on community‐level phenological patterns we compared patterns of first flowering dates (FFD) for herbaceous species at Konza Prairie, KS, USA with those from the colder Fargo, ND, USA area and from Chinnor, England, which has a less continental climate. Comparing patterns of FFD supports that Konza's flowering patterns are potentially influenced both by filtering species that flower early in the growing season and by phenological shifting. Konza species flowering dates were earlier in the spring and later in the fall compared to Fargo, but were not shifted compared to Chinnor, which had a unique suite of early‐flowering species. In all, comparing flowering phenology among three sites reveals that intraspecific responses to climate can generate phenological shifts that compress or stretch community‐level phenological patterns, while novel niches in phenological space can also alter community‐level patterns. Community flowering patterns related to climate suggest that climatic warming has the potential to further distribute flowering of the Konza flora over a longer period, but also could further open it to introductions of non‐native species that have evolved to flower early in the season.     

Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change

Although it is well established that butterfly richness is affected by climate and human factors (e.g. habitat disturbance and degradation) at different spatial scales, the drivers behind these changes vary greatly according to the geographical region and the ecology of the species concerned. It is essential that this variation be understood if trends in diversity are to be predicted with any degree of confidence under a scenario of global change. Here we examine patterns of butterfly species richness among groups differing in degree of habitat specialization, diet breadth and mobility in the north‐west Mediterranean Basin, a European hotspot for this taxon. We analyze a large number of butterfly communities and take into consideration the main potential drivers, that include climatic, geographic and resource variables, landscape structure and human environmental impact at different spatial scales. Our study shows that both climatic and anthropogenic factors play an important role in determining butterfly species richness in the north‐west Mediterranean Basin, but that their relative impact differs between specialist and generalist groups. At lower altitudes, water availability, a product of the interplay between temperature and rainfall, and negative effects of temperature appear as the most determinant factors. Maximum diversity was observed at mid‐altitudes, which reveals the importance from a conservation point of view of Mediterranean mountain ranges. The results suggest serious population declines in specialist species restricted to mountain areas as a result of climate warming in combination with habitat loss caused by the abandonment of grazing and mowing. They also suggest negative trends for generalist species due to an increase in aridity in combination with an increase in intensification of human land use in lowland areas. Such synergies are expected to lead to rapid declines in Mediterranean butterfly populations in the coming years, thereby posing a severe threat for the conservation of European biodiversity.     

Climate‐based empirical models show biased predictions of butterfly communities along environmental gradients

A better understanding of the factors that mould ecological community structure is required to accurately predict community composition and to anticipate threats to ecosystems due to global changes. We tested how well stacked climate‐based species distribution models (S‐SDMs) could predict butterfly communities in a mountain region. It has been suggested that climate is the main force driving butterfly distribution and community structure in mountain environments, and that, as a consequence, climate‐based S‐SDMs should yield unbiased predictions. In contrast to this expectation, at lower altitudes, climate‐based S‐SDMs overpredicted butterfly species richness at sites with low plant species richness and underpredicted species richness at sites with high plant species richness. According to two indices of composition accuracy, the Sorensen index and a matching coefficient considering both absences and presences, S‐SDMs were more accurate in plant‐rich grasslands. Butterflies display strong and often specialised trophic interactions with plants. At lower altitudes, where land use is more intense, considering climate alone without accounting for land use influences on grassland plant richness leads to erroneous predictions of butterfly presences and absences. In contrast, at higher altitudes, where climate is the main force filtering communities, there were fewer differences between observed and predicted butterfly richness. At high altitudes, even if stochastic processes decrease the accuracy of predictions of presence, climate‐based S‐SDMs are able to better filter out butterfly species that are unable to cope with severe climatic conditions, providing more accurate predictions of absences. Our results suggest that predictions should account for plants in disturbed habitats at lower altitudes but that stochastic processes and heterogeneity at high altitudes may limit prediction success of climate‐based S‐SDMs.     

Grasshopper community response to climatic change: variation along an elevational gradient

Background

The impacts of climate change on phenological responses of species and communities are well-documented; however, many such studies are correlational and so less effective at assessing the causal links between changes in climate and changes in phenology. Using grasshopper communities found along an elevational gradient, we present an ideal system along the Front Range of Colorado USA that provides a mechanistic link between climate and phenology.

Methodology/Principal Findings

This study utilizes past (1959–1960) and present (2006–2008) surveys of grasshopper communities and daily temperature records to quantify the relationship between amount and timing of warming across years and elevations, and grasshopper timing to adulthood. Grasshopper communities were surveyed at four sites, Chautauqua Mesa (1752 m), A1 (2195 m), B1 (2591 m), and C1 (3048 m), located in prairie, lower montane, upper montane, and subalpine life zones, respectively. Changes to earlier first appearance of adults depended on the degree to which a site warmed. The lowest site showed little warming and little phenological advancement. The next highest site (A1) warmed a small, but significant, amount and grasshopper species there showed inconsistent phenological advancements. The two highest sites warmed the most, and at these sites grasshoppers showed significant phenological advancements. At these sites, late-developing species showed the greatest advancements, a pattern that correlated with an increase in rate of late-season warming. The number of growing degree days (GDDs) associated with the time to adulthood for a species was unchanged across the past and present surveys, suggesting that phenological advancement depended on when a set number of GDDs is reached during a season.

Conclusions

Our analyses provide clear evidence that variation in amount and timing of warming over the growing season explains the vast majority of phenological variation in this system. Our results move past simple correlation and provide a stronger process-oriented and predictive framework for understanding community level phenological responses to climate change.     

Effect of regional climate warming on the phenology of butterflies in boreal forests in Manitoba, Canada

We examined the effect of regional climate warming on the phenology of butterfly species in boreal forest ecosystems in Manitoba, Canada. For the period 1971-2004, the mean monthly temperatures in January, September, and December increased significantly, as did the mean temperatures for several concurrent monthly periods. The mean annual temperature increased ≈ 0.05°C/yr over the study period. The annual number of frost-free days and degree-day accumulations increased as well. We measured the response of 19 common butterfly species to these temperature changes with the date of first appearance, week of peak abundance, and the length of flight period over the 33-yr period of 1972-2004. Although adult butterfly response was variable for spring and summer months, 13 of 19 species showed a significant (P < 0.05) increase in flight period extending longer into the autumn. Flight period extensions increased by 31.5 ± 13.9 (SD) d over the study period for 13 butterfly species significantly affected by the warming trend. The early autumn and winter months warmed significantly, and butterflies seem to be responding to this warming trend with a change in the length of certain life stages. Two species, Junonia coenia and Euphydryas phaeton, increased their northerly ranges by ≈ 150 and 70 km, respectively. Warmer autumns and winters may be providing opportunities for range extensions of more southerly butterfly species held at bay by past climatic conditions.     

Phenology responses of temperate butterflies to latitude depend on ecological traits

Global change influences species’ seasonal occurrence, or phenology. In cold‐adapted insects, the activity is expected to start earlier with a warming climate, but contradictory evidence exists, and the reactions may be linked to species‐specific traits. Using data from the GBIF database, we selected 105 single‐brooded Holarctic butterflies inhabiting broad latitudinal ranges. We regressed patterns of an adult flight against latitudes of the records, controlling for altitude and year effects. Species with delayed flight periods towards the high latitudes, or stable flight periods across latitudes, prevailed over those that advanced their flight towards the high latitudes. The responses corresponded with the species’ seasonality (flight of early season species was delayed and flight of summer species was advanced at high latitudes) and oceanic vs. continental climatic niches (delays in oceanic, stability in continental species). Future restructuring of butterfly seasonal patterns in high latitudes will reflect climatic niches, and hence the evolutionary history of participating species.     

Disproportional risk for habitat loss of high‐altitude endemic species under climate change

The expected upward shift of trees due to climate warming is supposed to be a major threat to range‐restricted high‐altitude species by shrinking the area of their suitable habitats. Our projections show that areas of endemism of five taxonomic groups (vascular plants, snails, spiders, butterflies, and beetles) in the Austrian Alps will, on average, experience a 77% habitat loss even under the weakest climate change scenario (+1.8 °C by 2100). The amount of habitat loss is positively related with the pooled endemic species richness (species from all five taxonomic groups) and with the richness of endemic vascular plants, snails, and beetles. Owing to limited postglacial migration, hotspots of high‐altitude endemics are situated in rather low peripheral mountain chains of the Alps, which have not been glaciated during the Pleistocene. There, tree line expansion disproportionally reduces habitats of high‐altitude species. Such legacies of climate history, which may aggravate extinction risks under future climate change have to be expected for many temperate mountain ranges.     

Tell me what you eat and I'll tell you when you fly: diet can predict phenological changes in response to climate change

Changes in phenology are correlated with climate change. However, we still struggle to understand the traits making species susceptible to climate change, and the implications of species' reactions for communities and food webs. Butterflies and moths are an ecologically important group that have shown pronounced phenological changes over the last decades. Tests using a > 150-year dataset from 566 European butterfly and moth species demonstrated that variation in phenological change was strongly related to traits describing plant-herbivore interactions (larval diet breadth, diet composition), and the life cycle. The results indicate that climate change related shifts in phenology are correlated with the seasonal availability and palatability of food plants. Lepidopterans feeding on herbaceous plants showed smaller shifts in flight periods but larger increases in voltinism than lepidopterans feeding on woody plants. Consequently, the effect of herbivorous lepidopterans may increase in herb-rich grassland ecosystems under warmer conditions, and not in forest ecosystems.