Climate Envelope Research Articles

Vegetation-rainfall coupling as an indicator of ecosystem state in a heterogeneous landscape

Coexisting vegetation types in tropical landscapes can respond in contrasting ways to rainfall, despite being in the same climatic envelope. Understanding such heterogeneity in vegetation-rainfall interactions is key for predicting how ecosystems might respond to future environmental changes. Here we test whether temporal coupling between vegetation greenness and rainfall is a good indicator of ecosystem state in the landscape. For this, we study a well-preserved landscape of the Brazilian Cerrado that is formed by mosaics of contrasting ecosystems, including savannas, dry forests and gallery forests. First, we correlate the time-series of rainfall and vegetation greenness to quantify their coupling for each vegetation type. We then compare vegetation-rainfall coupling with other state variables, such as local-scale vegetation structural and functional traits, as well as differences in environmental conditions in which these vegetation types exist. Coexisting vegetation types are set in contrasting local-scale environmental conditions and have distinct responsiveness to rainfall. Commonly used structural and functional state variables, such as tree cover and tree height, do not depict such marked differences between the vegetation types, particularly for gallery and dry forests. Dry forests have the strongest coupling and decrease their greenness during dry seasons, reflecting vegetation deciduousness on nutrient-richer soils. In contrast, gallery forests increase their greenness during the dry season, when direct radiation peaks, likely due to perennial access to groundwater. Savannas are less responsive to rainfall and have a more stable greenness throughout the year. Our findings suggest that heterogeneity in local abiotic conditions contribute to determining both vegetation distribution and ecosystem states in these tropical savanna landscapes. Changes in these conditions as a result of climate and land-use changes will likely alter the distribution of vegetation types in the future. Our functional metric may thus be useful for assessing future responses of tropical ecosystems to changes in precipitation.

Spatio-temporal shifts in British wild bees in response to changing climate.

Climate plays a major role in determining where species occur, and when they are active throughout the year. In the face of a changing climate, many species are shifting their ranges poleward. Many species are also shifting their emergence phenology. Wild bees in Great Britain are susceptible to changes in climatic conditions but little is known about historic or potential future spatio-temporal trends of many species. This study utilized a sliding window approach to assess the impacts of climate on bee emergence dates, estimating the best temperature window for predicting emergence dates for 88 species of wild bees. Using a 'middle-of-the-road' (RCP 4.5) and 'worst-case' (RCP 8.5) climate scenario for the period 2070-2079, predictions of future emergence dates were made. In general, the best predicting climate window occurred in the 0-3 months preceding emergence. Across the 40 species that showed a shift in emergence dates in response to a climate window, the mean advance was 13.4 days under RCP 4.5 and 24.9 days under RCP 8.5. Species distribution models (SDMs) were used to predict suitable climate envelopes under historic (1980-1989), current (2010-2019) and future (2070-2079 under RCP 4.5 and RCP 8.5 scenarios) climate conditions. These models predict that the climate envelope for 92% of studied species has increased since the 1980s, and for 97% and 93% of species under RCP 4.5 and RCP 8.5 respectively, this is predicted to continue, due to extension of the northern range boundary. While any range changes will be moderated by habitat availability, it highlights that Great Britain will likely experience northward shifts of bee populations in the future. By combining spatial and temporal trends, this work provides an important step towards informing conservation measures suitable for future climates, directing how interventions can be provided in the right place at the right time.

Climenv: Download, extract and visualise climatic and elevation data

AbstractUnderstanding the relationship between climate and vegetation requires climate data to be linked with ecological data, including habitat types and vegetation mapping. Our new R package climenv allows researchers to efficiently acquire, extract and visualise data sets that are commonly used by researchers to quantify the climatic envelope of vegetation. climenv offers integrated downloading and processing capabilities for three globally recognised data sets: WorldClim 2, CHELSA and the National Aeronautics and Space Administration's (NASA) Shuttle Radar Topography Mission (SRTM) elevation data. The package allows users to easily download and extract these data sets for single and multi‐geospatial polygon and point data sets, facilitating studies that explore the relationship between vegetation and climate. Furthermore, climenv allows users to plot traditional Holdridge life zone classification, Walter–Lieth climate diagrams and new customised plots, which combine aspects of both these systems with other biologically relevant climate variables. By enhancing the usability and flexibility of these data sets, climenv helps to explore the intricacies of the relationships between climate and vegetation. Our package is accessible from CRAN (https://CRAN.R‐project.org/package=climenv) or GitHub (https://github.com/jamestsakalos/climenv).

A maximum entropy approach to defining geographic bounds on growth and yield model usage

Growth and yield models are essential tools in modern forestry, especially for intensively managed loblolly pine plantations in the southeastern United States. While model developers often have a good idea of where these models should be used with respect to geographic location, determining geographic bounds for model usage can be daunting. Such bounds provide suitable areas where model predictions are likely to behave as expected or identify areas where models may do a poor job of characterizing the growth of a resource. In this research, we adapted a niche model methodology, commonly used to identify suitable spots for species occurrence (maximum entropy), to identify areas for using growth and yield models built from plots established in the Lower Coastal Plain and Piedmont/Upper Coastal Plain in the southeastern United States. The results from this analysis identify areas with similar climatic envelopes and soil properties to the areas where data was collected to fit these growth and yield models. These areas show notable overlap with the areas prescribed for use by the evaluated growth and yield models and support practitioners use of these models throughout these regions. Furthermore, this methodology can be applied to different forest models built using large regional extents as long as climatic and soil values are available for each site.

Surge-type glaciers in Kalaallit Nunaat (Greenland): distribution, temporal patterns and climatic controls

Abstract We present the first systematic inventory of surge-type glaciers for the whole of Greenland compiled from published datasets and multitemporal satellite images and digital elevation models. The inventory allows us to define the spatial and climatic distribution of surge-type glaciers and to analyse the timing of surges from 1985 to 2019. We identified 274 surge-type glaciers, an increase of 37% compared to previous work. Mapping surge-type glacier distribution by temperature and precipitation variables derived from ERA5-Land reanalysis data shows that the west and east clusters occur in well-defined climatic envelopes. Analysis of the timing of surge active phases during the periods ~1985 to 2000 (T1) and ~2000 to 2019 (T2) suggests that overall surge activity is similar in T1 and T2, but there appears to be a reduction in surging in the west cluster in T2. Our climate analysis shows a coincident increase in mean annual and mean winter air temperature between T1 and T2. We suggest that as glaciers thin under current warming, some surge-type glaciers in the west cluster may be being prevented from surging due to (1) their inability to build-up sufficient mass and (2) a switch from a polythermal to a largely cold-based thermal regime.

The conservation long game: Leiopelma species climate envelopes in New Zealand under a changing climate

The conservation long game: Leiopelma species climate envelopes in New Zealand under a changing climate

Characterization of the genetic composition and establishment of a core collection for the INERA Robusta coffee (Coffea canephora) field genebank from the Democratic Republic of Congo

Cultivation of Robusta coffee is likely to gain importance because of its high disease resistance and climate envelope. Robusta coffee genetic resources conserved in field genebanks can play an important role to further improve its cupping quality and other agronomic traits, but such Coffea canephora collections are limited and still poorly characterized. In this study, we characterized the genetic composition of the historically important but until recently neglected INERA Coffee Collection in Yangambi (the Democratic Republic of Congo). We used GBS to discover genome-wide genetic diversity, created and validated a novel multiplex amplicon sequencing (HiPlex) screening assay to genetically screen 730 coffee shrubs of the Yangambi Coffee Collection, grouped clonal material and delineated 263 accessions with unique genetic fingerprints. Comparison to reference material of three genetic origins revealed that the majority of the Yangambi accessions were assigned a ‘Lula’ cultivar origin, four accessions were assigned to Congolese subgroup A and nine accessions were most closely related to local wild accessions. About one-quarter of the accessions was likely derived from hybridization between these groups, which could result from seed-based propagation of the collection, breeding efforts, or natural cross-pollination. Parental analyses discovered eight preferentially used accessions, which may correspond to historically selected founders, or direct descendants thereof, whose seed material was once widely used to establish coffee plantations. Finally, two core collections were proposed using the maximization strategy (CC-I; 100 accessions) and genetic distance method (CC-X; 10 accessions). Our study demonstrates a method for the genetic characterization of Robusta coffee collections in general and contributes to the re-evaluation and exploration of the Robusta coffee genetic resources in the Democratic Republic of the Congo in particular.

Identifying conservation priority areas forNorth Americanbumble bee species inCanadaunder current and future climate scenarios

AbstractMany bumble bee species are declining globally from multiple threats including climate change. Identifying conservation priority areas with a changing climate will be important for conserving bumble bee species. Using systematic conservation planning, we identified priority areas for 44 bumble bee species in Canada under current and projected climates (year 2050). Conservation priority areas were identified as those that contained targeted amounts of each species predicted occurrence through climate envelope models, while minimizing the area cost of conserving the identified conservation priority areas. Conservation priority areas in the two periods were compared to established protected areas and land cover types to determine the area of current and future priority sites that are protected and the types of landscapes within priority areas. Notably, conservation priority areas were rarely within established protected areas. Priority areas were most often in croplands and grasslands, mainly within the mountain west, central and Southern Ontario, Northern Quebec, and Atlantic Canada under all climate scenarios. Conservation priority areas are predicted to increase in elevation and latitude with climate change. Our findings identify the most important regions in Canada for conserving bumble bee species under current and future climates including consistently selected future sites.

The climatic risk of Amazonian protected areas is driven by climate velocity until 2050.

Changes in species distribution in response to climate change might challenge the territorial boundaries of protected areas. Amazonia is one of the global regions most at risk of developing long distances between current and future analogous climates and the emergence of climate conditions without analogs in the past. As a result, species present within the network of Protected Areas (PAs) of Amazonia may be threatened throughout the 21st century. In this study, we investigated climate velocity based on future and past climate-analogs using forward and backward directions in the network of PAs of Amazonia, in order to assess the climatic risk of these areas to climate change and verify their effectiveness in maintaining the current climate conditions. Using current (1970-2000) and future (2041-2060) average annual air temperature and precipitation data with a resolution of 10 km, climate velocities across the entire Amazon biome and average climate velocities of PAs and Indigenous Lands (ILs) were evaluated. The results show that the effects of backward velocity will be greater than that of forward velocity in the Amazon biome. However, the PA network will be less exposed to backward velocity impacts than unprotected areas (UAs)-emphasizing the importance of these areas as a conservation tool. In contrast, for the forward velocity impacts, the PA network will be slightly more exposed than UAs-indicating that the current spatial arrangement of the PA network is still not the most suitable to minimize impacts of a possible climate redistribution. In addition, a large extent of no-analog climates for backward velocities was found in central Amazonia, indicating that high temperatures and changes in precipitation patterns in this region will surpass the historical variability of the entire biome, making it a potentially isolated and unsuitable climatic envelope for species in the future. Most of the no-analog climates are in PAs, however the climate risks in ILs should also be highlighted since they presented higher climate velocities than PAs in both metrics. Our projections contrast with the median latitudinal migration rate of 2 km/year observed in most ecosystems and taxonomic groups studied so far and suggest the need for median migration rates of 7.6 km/year. Thus, despite the important role of PAs and ILs as conservation tools, they are not immune to the effects of climate change and new management strategies, specific to each area and that allow adaptation to global changes, will be necessary.

Toward an extreme world: The hyper‐arid ecosystem as a natural model

AbstractThe forecasted increased frequency and intensity of extreme climatic events may strongly affect ecosystem structure and function in the future. It is unclear how ecosystems will function in the long run over a large spatial scale under a new extreme water cycle. This open question calls for a conceptual framework as a fundamental basis for theoretical and experimental exploration of ecosystem function on a large scale driven by an extreme climate envelope. To assess the problem on a large scale, we investigated hyper‐arid ecosystems (HAEs) as natural tangible models that already function under an extreme climatic envelope. Our new assertion is that if extreme climate change drives arid lands to function under alternate extreme conditions, then arid land ecosystems will function like an HAE as an alternative state, rather than progress to desertification. To support our assertion, we developed a conceptual framework of HAEs that includes a geo‐hydrological “abiotic engine” that drives HAE function by soil moisture diversity and plant functional groups. Based on this conceptual framework, we suggest incorporating two new hypotheses in climate change studies to advance our understanding of responses of large‐scale, water‐limited ecosystems: (1) Hydro‐climatic extremes in water‐limited ecosystems will reduce the degree of resource conservation by slope ecosystems due to reduction in plant cover and soil. The decreased ecosystem function on the slope will be compensated for by increasing the effect of the abiotic engine on the ephemeral stream, thus enhancing meta‐ecosystem functioning in the ephemeral stream. (2) In water‐limited ecosystems, climate change toward hydro‐climatic extremes will rescale the dominant hydro‐ecological processes of pulse–reserve, source–sink, and connectivity along the semiarid, arid, and HA gradients in two ways: (i) shrinking of both spatial and temporal dimensions; and (ii) shrinking in the temporal dimension and expanding in the spatial dimensions. The first rescaling trajectory is related to biodiversity–ecosystem function and the second to the abiotic engine processes.

Northward expansion of the southern-temperate podocarp forest during the early Eocene: Palynological evidence from the NE Tibetan Plateau (China)

The debated vegetation response to climate change can be investigated through palynological fossil records from past extreme climate conditions. In this context, the early Eocene (53.3 to 41.2 million years ago (Ma)) is often referred to as a model for a greenhouse Earth. In the Xining Basin, situated on the Northeastern Tibetan Plateau (NETP), this time interval is represented by an extensive and well-dated sedimentary sequence of evaporites and red mudstones. Here we focus on the palynological record of the Early Eocene Climatic Optimum (EECO; 53.3 to 49.1 Ma) and study the fossil gymnosperm pollen composition in these sediments. In addition, we also investigate the nearest living relatives (NLR) or botanical affinity of these genera and the paleobiogeographic implications of their occurrence in the Eocene of the NETP. To reach our objective, we complemented transmitted light microscopy with laser scanning- and electron microscopy techniques, to produce high-resolution images, and illustrate the morphological variation within fossil and extant gymnosperm pollen. Furthermore, a morphometric analysis was carried out to investigate the infra- and intrageneric variation of these and related taxa. To place the data in context we produced paleobiogeographic maps for Phyllocladidites and for other Podocarpaceae, based on data from a global fossil pollen data base, and compare these with modern records from GBIF. We also assessed the climatic envelope of the NLR. Our analyses confirm the presence of Phyllocladidites (NLR Phyllocladus, Podocarpaceae) and Podocarpidites (NLR Podocarpus, Podocarpaceae) in the EECO deposits in the Xining Basin. In addition, a comparative study based on literature suggests that Parcisporites is likely a younger synonym of Phyllocladidites. Our findings further suggest that the Phyllocladidites specimens are derived from a lineage that was much more diverse than previously thought, and which had a much larger biogeographical distribution during the EECO than at present. Based on the climatic envelope of the NLR, we suggest that the paleoclimatic conditions in the Xining Basin were warmer and more humid during the EECO. We conclude that phylloclade-type conifers typical of the southern-temperate podocarp forests, had a northward geographical expansion during the EECO, followed by extirpation.

Mechanistic movement models to predict geographic range expansions of ticks and tick-borne pathogens: Case studies with Ixodes scapularis and Amblyomma americanum in eastern North America

The geographic range of the blacklegged tick, Ixodes scapularis, is expanding northward from the United States into southern Canada, and studies suggest that the lone star tick, Amblyomma americanum, will follow suit. These tick species are vectors for many zoonotic pathogens, and their northward range expansion presents a serious threat to public health. Climate change (particularly increasing temperature) has been identified as an important driver permitting northward range expansion of blacklegged ticks, but the impacts of host movement, which is essential to tick dispersal into new climatically suitable regions, have received limited investigation. Here, a mechanistic movement model was applied to landscapes of eastern North America to explore 1) relationships between multiple ecological drivers and the speed of the northward invasion of blacklegged ticks infected with the causative agent of Lyme disease, Borrelia burgdorferi sensu stricto, and 2) its capacity to simulate the northward range expansion of infected blacklegged ticks and uninfected lone star ticks under theoretical scenarios of increasing temperature. Our results suggest that the attraction of migratory birds (long-distance tick dispersal hosts) to resource-rich areas during their spring migration and the mate-finding Allee effect in tick population dynamics are key drivers for the spread of infected blacklegged ticks. The modeled increases in temperature extended the climatically suitable areas of Canada for infected blacklegged ticks and uninfected lone star ticks towards higher latitudes by up to 31% and 1%, respectively, and with an average predicted speed of the range expansion reaching 61 km/year and 23 km/year, respectively. Differences in the projected spatial distribution patterns of these tick species were due to differences in climate envelopes of tick populations, as well as the availability and attractiveness of suitable habitats for migratory birds. Our results indicate that the northward invasion process of lone star ticks is primarily driven by local dispersal of resident terrestrial hosts, whereas that of blacklegged ticks is governed by long-distance migratory bird dispersal. The results also suggest that mechanistic movement models provide a powerful approach for predicting tick-borne disease risk patterns under complex scenarios of climate, socioeconomic and land use/land cover changes.

Insights from the first winter weather observations near Mount Everest's summit

Insights from the first winter weather observations near Mount Everest's summit

Climatic envelopes of the genus Lacerta Linnaeus, 1758 in Türkiye: an application of ecological niche modeling.

Six species belonging to the genus Lacerta live in Türkiye. In this study, both present and future potential distribution maps were created based on occurrence data and climatic variables for these six species. Two scenarios for future projections (shared socioeconomic pathways, SSPs,: 245 and 585) and two timeframes (2041-2060 and 2081-2100) were used. The present and future potential distributions of these species were compared. As a result, it was predicted that the distribution ranges in the six species will expand in the future, and this expansion has revealed new environments.

Formation of the Holarctic Fauna: Dated molecular phylogenetic and biogeographic insights from the Quedius-lineage of Ground-Dwelling Rove Beetles (Coleoptera, Staphylinidae)

Although the Holarctic fauna has been explored for centuries, many questions on its formation are still unanswered. For example, i) what was the impact of the uplift of the Himalaya and Tibetan Plateau?, ii) what were the timings and climate of the faunal bridges connecting the Nearctic and Palearctic regions?, and iii) how did insect lineages respond to the late Paleogene global cooling and regional aridification? To answer these, we developed a phylogenetic dataset of 1229 nuclear loci for a total of 222 species of rove beetles (Staphylinidae) with emphasis in the tribe Quediini, especially Quedius-lineage and its subclade Quedius sensu stricto. Using eight fossils for calibrating molecular clock, we estimated divergence times and then analysed in BioGeoBEARS paleodistributions of the most recent common ancestor for each target lineage. For each species we generated climatic envelopes of the temperature and precipitation and mapped them across the phylogeny to explore evolutionary shifts. Our results suggest that the warm and humid Himalaya and Tibetan Plateau acted as an evolutionary cradle for the Quedius-lineage originating during the Oligocene from where, in the Early Miocene, the ancestor of the Quedius s. str. dispersed into the West Palearctic. With the climate cooling from the Mid Miocene onwards, new lineages within Quedius s. str. emerged and gradually expanded distributions across the Palearctic. In Late Miocene, a member of the group dispersed to the Nearctic region via Beringia before the closure of this land bridge 5.3 Ma. Paleogene global cooling and regional aridification largely shaped the current biogeographic pattern for Quedius s. str. species, many of them originating during the Pliocene and shifting or contracting their ranges during Pleistocene.

Climate-related range shifts in Arctic-breeding shorebirds.

To test whether the occupancy of shorebirds has changed in the eastern Canadian Arctic, and whether these changes could indicate that shorebird distributions are shifting in response to long-term climate change. Foxe Basin and Rasmussen Lowlands, Nunavut, Canada. We used a unique set of observations, made 25 years apart, using general linear models to test if there was a relationship between changes in shorebird species' occupancy and their species temperature Index, a simple version of a species climate envelope. Changes in occupancy and density varied widely across species, with some increasing and some decreasing. This is despite that overall population trends are known to be negative for all of these species based on surveys during migration. The changes in occupancy that we observed were positively related to the species temperature index, such that the warmer-breeding species appear to be moving into these regions, while colder-breeding species appear to be shifting out of the regions, likely northward. Our results suggest that we should be concerned about declining breeding habitat availability for bird species whose current breeding ranges are centered on higher and colder latitudes.

Climate Change and Animal Movement Integration in the Environmental Niche Model

Changing precipitation and temperature patterns due to climate change, shift ecological niches which pose a challenge for species. Furthermore, it is still unclear that if climate change faster than the speed of the species to move to more suitable environments. Climate Envelope models (GEMs) are used extensively in this matter to predict species geographical distribution. In this study, climate data and animal movement strategies integrated into the environmental niche model to analyze the successes of the species that have different movement strategies under the changing climate conditions. Four different movement strategies are formulized; lazy knowledge strategy (LKS), lazy no knowledge strategy (LNKS), proactive knowledge (PKS) and proactive no knowledge strategy (PNKS). In this study, mean annual temperature and annual precipitation data gathered for the RCP8.5 scenario from the HadGEM2-ES GCM model at a 10-minute resolution. Results show that the PKS has the highest survival rates which makes it the most successful strategy. Furthermore, the species with knowledge had performed higher survival rates than the species without knowledge due to their skill to find the most suitable cells around them. One-Way Anova test confirmed that there are significant differences between the strategies. Moreover, the analysis suggests that the species with knowledge of their environment have the most successful strategies for facing the climate change.

The dynamics of a changing Lutz spruce (Picea×lutzii) hybrid zone on the Kenai Peninsula, Alaska

We investigated the genetic makeup of Lutz spruce, a natural hybrid between white and Sitka spruce on the Kenai Peninsula, Alaska. Microsatellites indicate 72% of individuals sampled had predominantly white spruce ancestry, whereas 14% had predominantly Sitka spruce ancestry; some individuals classified as white spruce had Sitka spruce plastid genotypes. As Picea mitochondria are maternally inherited and plastids are paternally inherited, it appears that white spruce was the ancestral seed parent of nearly all spruce on the western peninsula, whereas Sitka spruce alleles originated from pollen. Pollen records show that white spruce colonized the western peninsula ∼8500 YBP from glacial refugium, whereas Sitka spruce arrived on the eastern peninsula ∼4000 YBP after migrating up the Pacific coast. Our data suggest that Sitka spruce colonization west of the Kenai Mountains may have occurred not via seed dispersal but by long distance transport of wind-borne pollen and subsequent hybridization with established white spruce populations. Hybridization was an important mechanism that allowed Sitka spruce to expand the leading edge of its range in response to historical climate change. As the climate continues to warm, climate envelope modeling suggests Lutz spruce may ultimately displace white spruce on the western peninsula even as Sitka spruce is constrained to the eastern peninsula where it will continue to hybridize.

Optimism for the Future Use of Environmental DNA (eDNA) to Study Global Climate Change

Optimism for the Future Use of Environmental <scp>DNA</scp> (<scp>eDNA</scp>) to Study Global Climate Change

Speed of extirpation of the huemul in the history of human occupation in Patagonia

Context The Patagonian huemul, an endangered Odocoilinedeer, has an estimated 350–500 individuals remaining in Argentina. Today’s population size, representing a numerical reduction of &gt;99% of original estimates, is fragmented into small groups along ~2000 km of Andean mountains. The species’ numbers were heavily reduced by past overexploitation and they disappeared in areas of high anthropogenic activity, predominantly the fertile valley bottoms. Aims This research delineates the current potential distribution of Patagonian huemul by using climatic indicators, topographic and vegetational proxies, and anthropogenic pressure, to determine the relevance of the climatic envelope on current distribution. Methods Occurrence records (latitude and longitude) were compiled (n = 159) by consulting the literature. Twenty environmental variables were used (WorldClim database) and two other representative environmental variables (normalised difference vegetation index (NDVI) and enhanced vegetation index (EVI)) were added to test their predictive power. We added the human footprint index (HFP) as a variable to control for model bias. Using the maximum entropy algorithm (MaxEnt), we modelled the species’ potential distribution. We designated the historical distribution as area M. Additionally, we calculated three areas of distribution: current, historical and potential. Finally, we calculated distributional retraction of the species and area lost per year. Key results The model showed good predictive power (AUCTest = 0.764 ± 0.091). However, low values were obtained for AUCtrain and AUCprom for the different predictor scenarios. Although the model shows the interaction among several climatic, environmental, and topographic variables, the human footprint index (39.9%) was the variable that most influenced the current potential distribution of this species. Conclusion Our model shows that most of Patagonia’s surface is climatically suitable for huemul. This suggests that the causes of distributional retraction are not related to limitations imposed by the climate envelope, but rather concur with recent research showing impact owing to the species’ behavioural response to anthropogenic activity. Implications Current populations are small, fragmented, and confined to poor-quality sites. Although the species is currently found mainly within protected areas, management actions must be initiated that promote innovative strategies in unprotected areas, as well as high-value habitats, particularly as protected areas contain limited fertile lower-valley habitats.