Landscape-scale Heterogeneity Research Articles

Achieving structural heterogeneity and high multi-taxon biodiversity in managed forest ecosystems: a European review

AbstractThe European Biodiversity Strategy has set the key goal to maintain and promote biodiversity. Managed forests here play a key role, as they are among the most diverse ecosystems. To create biodiversity rich managed forest landscapes, we need a deep understanding on how management affects forest structure and subsequently habitat quality on the local and landscape level. However, to date a confusingly large amount of different terms for various management systems exist and it remains unclear how forest structure and composition affected by specific management systems affect biodiversity. Here, we first aim to clarify forest management systems terminology. Second, we link existing management systems with forest structure and review at European level how these structures affect local α-, as well as landscape-scale β- and γ-biodiversity. We found that research and derived management guidelines have a strong focus on local forest stand heterogeneity and related α-diversity, while ignoring the importance of landscape-scale heterogeneity and therefore β- and γ-diversity. Instead of promoting one management system as an all-in-one-solution, a diversity of different forest management systems seems the most promising way to create biodiversity rich forest landscapes. We finally discuss how a combination of different management systems might help to create structurally rich forest habitats and landscapes, simulating different successional stages and promoting species rich communities.

Sustainability implications of carbon dynamics on the avocado frontier

Carbon storage is examined along the avocado frontier in the state of Michoacán, Mexico, where native pine-oak forest is being rapidly lost for the sake of avocado production expansion. Carbon pools for aboveground biomass carbon and for belowground soil carbon were compared across avocado orchards and neighboring pine-oak forests. While aboveground carbon was found to be significantly higher in forests when compared to orchards, there was no significant difference found in soil carbon stocks between the two land uses. Mixed effects regression models were used to parse out the landscape scale heterogeneity that is affecting soil carbon beyond land cover alone. Temperature, precipitation, and fragmentation variables were all found to significantly impact soil carbon across the landscape studied. Furthermore, there is evidence of over-fertilization within orchards, particularly of phosphorus, which may be temporarily subsidizing soil carbon loss. Sustainability considerations, therefore, must consider land use decisions in relation to other soil nutrients, beyond just carbon, to comprehensively address sustainability concerns arising from the impacts of avocado production.

Landscape-scale connectivity and fragment size determine species composition of grassland fragments

As a consequence of agricultural intensification and habitat fragmentation since the mid-20th century, biological diversity has declined considerably throughout the world, particularly in Europe. We assessed how habitat and landscape-scale heterogeneity, such as variation in fragment size (small vs. large) and landscape configuration (measured as connectivity index), affect plant and arthropod diversity. We focused on arthropods with different feeding behaviour and mobility, spiders (predators, moderate dispersal), true bugs (mainly herbivores and omnivores with moderate dispersal), wild bees (pollinators with good dispersal abilities), and wasps (pollinators, omnivores with good dispersal abilities). We studied 60 dry grassland fragments in the same region (Hungarian Great Plain); 30 fragments were represented by the grassland component of forest-steppe stands, and 30 were situated on burial mounds (kurgans). Forest-steppes are mosaics of dry grasslands with small forests in a matrix of plantation forests. Kurgans are ancient burial mounds with moderately disturbed grasslands surrounded by agricultural fields. The size of fragments ranged between 0.16–6.88 ha (small: 0.16–0.48 ha, large: 0.93–6.88 ha) for forest-steppes and 0.01–0.44 ha (small: 0.01–0.10 ha and large: 0.20–0.44 ha) for kurgans. Fragments also represented an isolation gradient from almost cleared and homogenous landscapes, to landscapes with relatively high compositional heterogeneity. Fragment size, connectivity, and their interaction affected specialist and generalist species abundances of forest-steppes and kurgans. Large fragments had higher species richness of ground-dwelling spiders, and the effect of connectivity was more strongly positive for specialist arthropods and more strongly negative for generalists in large than in small fragments. However, we also found a strong positive impact of connectivity for generalist plants in small kurgans in contrast to larger ones. We conclude that besides the well-known effect of enhancing habitat quality, increasing connectivity between fragments by restoring natural and semi-natural habitat patches would help to maintain grassland biodiversity.

Heterogeneity in biological assemblages and exposure in chemical risk assessment: Exploring capabilities and challenges in methodology with two landscape-scale case studies

Chemical exposure concentrations and the composition of ecological receptors (e.g., species) vary in space and time, resulting in landscape-scale (e.g. catchment) heterogeneity. Current regulatory, prospective chemical risk assessment frameworks do not directly address this heterogeneity because they assume that reasonably worst-case chemical exposure concentrations co-occur (spatially and temporally) with biological species that are the most sensitive to the chemical’s toxicity. Whilst current approaches may parameterise fate models with site-specific data and aim to be protective, a more precise understanding of when and where chemical exposure and species sensitivity co-occur enables risk assessments to be better tailored and applied mitigation more efficient. We use two aquatic case studies covering different spatial and temporal resolution to explore how geo-referenced data and spatial tools might be used to account for landscape heterogeneity of chemical exposure and ecological assemblages in prospective risk assessment. Each case study followed a stepwise approach: i) estimate and establish spatial chemical exposure distributions using local environmental information and environmental fate models; ii) derive toxicity thresholds for different taxonomic groups and determine geo-referenced distributions of exposure-toxicity ratios (i.e., potential risk); iii) overlay risk data with the ecological status of biomonitoring sites to determine if relationships exist. We focus on demonstrating whether the integration of relevant data and potential approaches is feasible rather than making comprehensive and refined risk assessments of specific chemicals. The case studies indicate that geo-referenced predicted environmental concentration estimations can be achieved with available data, models and tools but establishing the distribution of species assemblages is reliant on the availability of a few sources of biomonitoring data and tools. Linking large sets of geo-referenced exposure and biomonitoring data is feasible but assessment of risk will often be limited by the availability of ecotoxicity data. The studies highlight the important influence that choices for aggregating data and for the selection of statistical metrics have on assessing and interpreting risk at different spatial scales and patterns of distribution within the landscape. Finally, we discuss approaches and development needs that could help to address environmental heterogeneity in chemical risk assessment.

Simulated treatment effects on bird communities inform landscape-scale dry conifer forest management.

Human land use and climate change have increased forest density and wildfire risk in dry conifer forests of western North America, threatening various ecosystem services, including habitat for wildlife. Government policy supports active management to restore historical structure and ecological function. Information on potential contributions of restoration to wildlife habitat can allow assessment of tradeoffs with other ecological benefits when prioritizing treatments. We predicted avian responses to simulated treatments representing alternative scenarios to inform landscape‐scale forest management planning along the Colorado Front Range. We used data from the Integrated Monitoring in Bird Conservation Regions program to inform a hierarchical multispecies occupancy model relating species occupancy and richness with canopy cover at two spatial scales. We then simulated changes in canopy cover (remotely sensed in 2018) under three alternative scenarios, (1) a “fuels reduction” scenario representing landscape‐wide 30% reduction in canopy cover, (2) a “restoration” scenario representing more nuanced, spatially variable treatments targeting historical conditions, and (3) a reference, no‐change scenario. Model predictions showed areas of potential gains and losses for species richness, richness of ponderosa pine forest habitat specialists, and the ratio of specialists to generalists at two (1 km2 and 250 m2) spatial scales. Under both fuels reduction and restoration scenarios, we projected greater gains than losses for species richness. Surprisingly, despite restoration more explicitly targeting ecologically relevant historical conditions, fuels reduction benefited bird species richness over a greater spatial extent than restoration, particularly in the lower montane life zone. These benefits reflected generally positive species associations with moderate canopy cover promoted more consistently under the fuels reduction scenario. In practice, contemporary forest management is likely to lie somewhere between the fuels reduction and restoration scenarios represented here. Therefore, our results inform where and how active forest management can best support avian diversity. Although our study raises questions regarding the value of including landscape‐scale heterogeneity as a management objective, we do not question the value of targeting finer scale heterogeneity (i.e., stand and treatment level). Rather, our results combined with those from previous work clarify the scale at which targeting structural heterogeneity and historical reference conditions can promote particular ecosystem services.

Leaf traits and canopy structure together explain canopy functional diversity: an airborne remote sensing approach.

Plant functional diversity is strongly connected to photosynthetic carbon assimilation in terrestrial ecosystems. However, many of the plant functional traits that regulate photosynthetic capacity, including foliar nitrogen concentration and leaf mass per area, vary significantly between and within plant functional types and vertically through forest canopies, resulting in considerable landscape-scale heterogeneity in three dimensions. Hyperspectral imagery has been used extensively to quantify functional traits across a range of ecosystems but is generally limited to providing information for top of canopy leaves only. On the other hand, lidar data can be used to retrieve the vertical structure of forest canopies. Because these data are rarely collected at the same time, there are unanswered questions about the effect of forest structure on the three -dimensional spatial patterns of functional traits across ecosystems. In the United States, the National Ecological Observatory Network's Airborne Observation Platform (NEON AOP) provides an opportunity to address this structure-function relationship by collecting lidar and hyperspectral data together across a variety of ecoregions. With a fusion of hyperspectral and lidar data from the NEON AOP and field-collected foliar trait data, we assessed the impacts of forest structure on spatial patterns of N. In addition, we examine the influence of abiotic gradients and management regimes on top-of-canopy percent N and total canopy N (i.e., the total amount of N [g/m2 ] within a forest canopy) at a NEON site consisting of a mosaic of open longleaf pine and dense broadleaf deciduous forests. Our resulting maps suggest that, in contrast to top of canopy values, total canopy N variation is dampened across this landscape resulting in relatively homogeneous spatial patterns. At the same time, we found that leaf functional diversity and canopy structural diversity showed distinct dendritic patterns related to the spatial distribution of plant functional types.

Steppe Marmot (Marmota bobak) as ecosystem engineer in arid steppes

Burrow-dwelling rodents are often considered ecosystem engineer species in arid environments. They create distinct habitat patches by building burrows: they move large amounts of soil, mix soil layers and change soil properties locally. Our aim was to explore the role of Steppe Marmot as an ecosystem engineer in shaping the plant species composition and diversity of steppes. First, we made a literature search to gather information on the ecosystem engineering effect of the species. Second, in a case study, we compared the vegetation of marmot burrows with the surrounding intact steppes in North-Kazakhstan to identify differences in species composition and plant functional groups. Vegetation of the burrows was structurally and compositionally different from the intact steppe vegetation. Burrows were characterised by lower total vegetation cover, higher cover of annuals and lower cover of perennial grasses compared to the intact steppe. We found an increased cover of ruderal species on the burrows, but also several specialist species, such as Agropyron cristatum, Anabasis salsa, Kochia prostrata and Petrosimonia spp. were confined to the burrow vegetation. Our results suggest that marmot burrows increase the landscape-scale heterogeneity of the steppe vegetation and could act as stepping stones for the dispersal of several steppe-specialist species.

Environmental drivers of the forest regeneration in temperate mixed forests

As modern silviculture in natural forests is based on natural regeneration, finding the most important drivers of regeneration is crucial for forestry as well as conservation. We explored the relationship between numerous environmental and land use history variables and the species richness, cover and composition of the regeneration layer, and also the cover of the dominant species of the regeneration (sessile oak, hornbeam and beech) in coniferous-deciduous mixed forests. We identified the key factors which forest management can influence to support the regeneration of mixedwoods.Thirty-four stands were sampled, representing different tree species combinations and stand structures. We used redundancy analysis to explore the effects of the explanatory variables on the regeneration’s species composition, and general linear modelling to examine their effects on its species richness and cover.The most important drivers of species composition were tree species richness, the amount of relative diffuse light, the proportion of beech in the overstory, and the heterogeneity of the diameter of trees. The cover of the regeneration layer was positively related to the density of large trees and to the amount of relative diffuse light. Its species richness was most strongly influenced by light and tree species richness. For the cover of a particular species in the regeneration, the proportion of the conspecific species in the overstory was determinant for every species, but other, various drivers also played a role in the case of the different species.According to our results, the community variables of the regeneration are mainly driven by the characteristics of the current forest stands, thus they are strongly influenced by management. Compositional heterogeneity of the overstory, various tree size distribution and the presence of large trees play key roles in the maintenance of a heterogeneous regeneration layer. The shelterwood forestry system is partially capable of providing these conditions, but continuous cover forestry is much more suitable to achieve them. Besides the stand structural variables, among the drivers of the individual species, various variables of forest site, landscape and land use history also occurred. Therefore, we conclude that maintaining the landscape-scale heterogeneity of forest types and management systems may promote the coexistence of various species in the region.

Local and landscape-scale heterogeneity shape spotted wing drosophila (Drosophila suzukii) activity and natural enemy abundance: Implications for trophic interactions

Like natural enemies, invasive polyphagous pests may respond positively to local and landscape-scale diversity, making them more serious challenges to sustainable management than specialists. Yet the current view of landscape effects on pests rarely considers perennial systems or polyphagous pests. We investigated Drosophila suzukii (SWD) and natural enemy distributions and trophic interactions in blueberry systems of the southeast US in relation to local management and landscape structure. Fields were nested in 1-km landscapes along gradients of crop and forest heterogeneity (landscape compositional diversity) and field border and crop border lengths (landscape configurational diversity). Yeast-sugar bait traps were used to collect SWD and canopy suction sampling for natural enemies. Management practices at the local scale did not influence SWD abundance, but natural enemy abundance was higher in organic systems and fields with vegetation present between rows. Landscape configurational diversity and greater proportion of non-cropping habitat promoted higher SWD populations. Natural enemy communities were composed of highly generalized species, and contrary to our predictions, we observed lower abundances in landscapes with higher proportion of non-cropping area. Few predators were detected to have fed on SWD, and communities were dominated by predators with low specialization on prey in blueberry. Our results indicate that local and landscape features in blueberry landscapes have differential effects on a polyphagous pest and generalist natural enemies. Our initial analysis of predator diets, using a metabarcoding approach, suggests spatial structure in specialization on prey with high diversity of prey in predators collected in forested margins.

Topography shapes the structure, composition and function of tropical forest landscapes.

Topography is a key driver of tropical forest structure and composition, as it constrains local nutrient and hydraulic conditions within which trees grow. Yet, we do not fully understand how changes in forest physiognomy driven by topography impact other emergent properties of forests, such as their aboveground carbon density (ACD). Working in Borneo – at a site where 70‐m‐tall forests in alluvial valleys rapidly transition to stunted heath forests on nutrient‐depleted dip slopes – we combined field data with airborne laser scanning and hyperspectral imaging to characterise how topography shapes the vertical structure, wood density, diversity and ACD of nearly 15 km2 of old‐growth forest. We found that subtle differences in elevation – which control soil chemistry and hydrology – profoundly influenced the structure, composition and diversity of the canopy. Capturing these processes was critical to explaining landscape‐scale heterogeneity in ACD, highlighting how emerging remote sensing technologies can provide new insights into long‐standing ecological questions.

How landscape organization and scale shape catchment hydrology and biogeochemistry: insights from a long‐term catchment study

Catchment science plays a critical role in the protection of water resources in the face of ongoing changes in climate, long‐range transport of air pollutants, and land use. Addressing these challenges, however, requires improved understanding of how, when, and where changes in water quantity and quality occur within river networks. To reach these goals, we must recognize how different catchment features are organized to regulate surface chemistry at multiple scales, from processes controlling headwaters, to the downstream mixing of water from multiple landscape sources and deep aquifers. Here we synthesize 30‐years of hydrological and biogeochemical research from the Krycklan catchment study (KCS) in northern Sweden to demonstrate the benefits of coupling long‐term monitoring with multi‐scale research to advance our understanding of catchment functioning across space and time. We show that the regulation of hydrological and biogeochemical patterns in the KCS can be decomposed into four, hierarchically structured landscape features that include: (1) transmissivity and reactivity of dominant source layers within riparian soils, (2) spatial arrangement of groundwater input zones that govern water and solute fluxes at reach‐ to segment‐scales, (3) landscape scale heterogeneity (forests, mires, and lakes) that generates unique biogeochemical signals downstream, and (4) broad‐scale mixing of surface streams with deep groundwater contributions. While this set of features are perhaps specific to the study region, analogous hierarchical controls are likely to be widespread. Resolving these scale dependent processes is important for predicting how, when, and where different environmental changes may influence patterns of surface water chemistry within river networks. WIREs Water 2018, 5:e1265. doi: 10.1002/wat2.1265This article is categorized under: Science of Water > Hydrological Processes Science of Water > Water and Environmental Change Science of Water > Methods

Landscape scale heterogeneity in the East Turkana ecosystem during the Okote Member (1.56–1.38 Ma)

Placing the biological adaptations of Pleistocene hominins within a well-resolved ecological framework has been a longstanding goal of paleoanthropology. This effort, however, has been challenging due to the discontinuous nature of paleoecological data spanning many important periods in hominin evolution. Sediments from the Upper Burgi (1.98–1.87 Ma), KBS (1.87–1.56 Ma) and Okote (1.56–1.38 Ma) members of the Koobi Fora Formation at East Turkana in northern Kenya document an important time interval in the evolutionary history of the hominin genera Homo and Paranthropus. Although much attention has been paid to Upper Burgi and KBS member deposits, far less is known regarding the East Turkana paleoecosystem during Okote Member times. This study pairs spatially-resolved faunal abundance data with stable isotope geochemistry from mammalian enamel to investigate landscape-scale ecosystem variability during Okote Member times. We find that during this period 1) taxa within the East Turkana large mammal community were distributed heterogeneously across space, 2) the abundance of C3 and C4 vegetation varied between East Turkana subregions, and 3) the Karari subregion, an area with abundant evidence of hominin stone tool manufacture, had significantly more C3 vegetation than regions closer to the central axis of the Turkana Basin (i.e., Ileret and Koobi Fora). These findings indicate that the East Turkana paleoecosystem during the Okote Member was highly variable across space and provided a complex adaptive landscape for Pleistocene hominins.

Environmental filtering, local site factors and landscape context drive changes in functional trait composition during tropical forest succession

Second-growth forests provide an important avenue for at least partially recuperating biodiversity and ecosystem services in tropical regions. Yet, factors affecting changes in species and functional trait composition of trees during forest succession remain poorly understood. The environmental filtering hypothesis of community assembly predicts gradual successional changes in functional trait composition. However, differences in landscape conditions and local site factors can lead to idiosyncratic or divergent changes in species and functional composition.We examine community assembly patterns of canopy trees and palms during natural regeneration using functional trait measurements and 18 years of annual vegetation dynamics data from six second-growth and two old-growth forest stands in northeast Costa Rica. We measured eight functional traits on 94 species, which cumulatively accounted for at least 80% of the abundance in each plot. Old-growth specialists had significantly lower leaf nitrogen and phosphorus content and significantly higher wood specific gravity than second-growth specialists. At the community level, specific leaf area, leaf nitrogen content, leaf dry matter content, and leaf toughness showed directional trends from more acquisitive trait values towards more conservative trait values. The landscape context was also an important driver of the successional trends within and among plots, as most functional trait patterns were influenced by the location of the plots within the matrix. Furthermore, the differential abundance of palms among stands strongly influenced functional trait differences among plots. After accounting for the effects of stand basal area and stand location, random plot effects explained an additional 45–90% of the variation in community-weighted mean functional traits. Local and landscape scale heterogeneity often influenced functional trait variation more strongly than stand basal area, suggesting that both stochasticity and environmental filtering influence species and functional turnover during succession.

Vegetation history, recent dynamics and future prospects of a Hungarian sandy forest-steppe reserve: forest-grassland relations, tree species composition and size-class distribution

Pannonian forest-steppes host a high number of endemic species and contribute to landscape-scale heterogeneity. Alterations in the proportion of forests and grasslands due to changes in land-use practice and climatic parameters can have serious nature conservation consequences. Hypotheses about forest-steppe dynamics have rarely been verified by detailed analyses, especially for the sandy forest-steppes. We integrated historical analysis, aerial photo interpretation and field investigation to determine how vegetation of a sandy forest-steppe has changed, how current dynamical processes operate and how native and exotic tree species regenerate under present conditions. The vegetation of the study area before the onset of major anthropogenic environmental transformations in the Carpathian Basin may have been a mosaic of forested and unforested patches. However, there is strong evidence that after heavy deforestation, the region was almost completely treeless between the 15th and the 19th centuries. Forest cover was able to recover by the 1800s but the lack of forested areas in the region for centuries explains why forest patches are still poor in species. Grasslands, which existed continuously, are more diverse, supporting several rare and endemic species. From 1953 till 2013, 72.45% of the area proved to be stable, but 27.55% showed clear dynamical character, changing either from forest to grassland, or vice versa. Thus, cyclic dynamics can occur in sandy forest-steppes. We found that forest patches of different size, differently exposed edges and grasslands provide different habitats for the tree species. Exotic species were present in large numbers, probably due to the small size of the reserve and the lack of a buffer zone.

An Integrative Analysis of the Dynamics of Landscape- and Local-Scale Colonization of Mediterranean Woodlands by Pinus halepensis

Afforestation efforts have resulted in extensive plantations of either native or non-native conifers, which in many regions has led to the spread of those conifers into surrounding natural vegetation. This process of species colonization can trigger profound changes in both community dynamics and ecosystem processes. Our study disentangled the complexity of a process of colonization in a heterogeneous landscape into a simple set of rules. We analyzed the factors that control the colonization of natural woodland ecosystems by Pinus halepensis dispersing from plantations in the Mediterranean region of Israel. We developed maximum-likelihood models to explain the densities of P. halepensis colonizing natural woodlands. Our models unravel how P. halepensis colonization is controlled by factors that determine colonization pressure by dispersing seeds and by factors that control resistance to colonization of the natural ecosystems. Our models show that the combination of different seed arrival processes from local, landscape, and regional scales determine pine establishment potential, but the relative importance of each component varied according to seed source distribution. Habitat resistance, determined by abiotic and biotic conditions, was as important as propagule input in determining the density of pine colonization. Thus, despite the fact that pine propagules disperse throughout the landscape, habitat heterogeneity within the natural ecosystems generates significant variation in the actual densities of colonized pine. Our approach provides quantitative measures of how processes at different spatial scales affect the distribution and densities of colonizing species, and a basis for projection of expected distributions. Variation in colonization rates, due to landscape-scale heterogeneity in both colonization pressure and resistance to colonization, can be expected to produce a diversity of new ecosystems. This work provides a template for understanding species colonization processes, especially in light of anthropogenic impacts, and predicting future transformation of natural ecosystems by species invasion.

Evaluation of Two Energy Balance Closure Parametrizations

A general lack of energy balance closure indicates that tower-based eddy-covariance (EC) measurements underestimate turbulent heat fluxes, which calls for robust correction schemes. Two parametrization approaches that can be found in the literature were tested using data from the Canadian Twin Otter research aircraft and from tower-based measurements of the German Terrestrial Environmental Observatories (TERENO) programme. Our analysis shows that the approach of Huang et al. (Boundary-Layer Meteorol 127:273–292, 2008), based on large-eddy simulation, is not applicable to typical near-surface flux measurements because it was developed for heights above the surface layer and over homogeneous terrain. The biggest shortcoming of this parametrization is that the grid resolution of the model was too coarse so that the surface layer, where EC measurements are usually made, is not properly resolved. The empirical approach of Panin and Bernhofer (Izvestiya Atmos Oceanic Phys 44:701–716, 2008) considers landscape-level roughness heterogeneities that induce secondary circulations and at least gives a qualitative estimate of the energy balance closure. However, it does not consider any feature of landscape-scale heterogeneity other than surface roughness, such as surface temperature, surface moisture or topography. The failures of both approaches might indicate that the influence of mesoscale structures is not a sufficient explanation for the energy balance closure problem. However, our analysis of different wind-direction sectors shows that the upwind landscape-scale heterogeneity indeed influences the energy balance closure determined from tower flux data. We also analyzed the aircraft measurements with respect to the partitioning of the “missing energy” between sensible and latent heat fluxes and we could confirm the assumption of scalar similarity only for Bowen ratios $$\approx $$ 1.

Interdisciplinary historical vegetation mapping for ecological restoration in Galapagos

Improving our knowledge of pre-anthropogenic landscapes is vital for understanding landscape-scale heterogeneity and for setting goals and objectives for ecological restoration. This is especially important in highly modified landscapes that contain few remnants of pre-impact ecosystems. This study aims to develop new methodology to improve understanding of historical vegetation, using the now-degraded inhabited highlands of the Galapagos Islands as a case study. Our multidisciplinary approach innovatively combines data from interviews with residents who were familiar with the vegetation before most degradation occurred with the more traditional sources of historical aerial photography and information from early explorer and scientist reports. We reconstruct historical vegetation across the landscape by mapping it in the year 1960 and discussing this map in the historical context of anthropogenic change. Our results confirm published vegetation types but also define some other types not previously described, and suggest much greater spatial, temporal and structural heterogeneity than commonly understood. This result can be used by Galapagos land managers to better match species assemblages with sites and plan restoration actions that will maximise resilience against the ongoing and future threats of climate change and species invasions. Our methodology can be applied in extensive areas of the world where the majority of anthropogenic disturbance to natural ecosystems has been within the past 60 years.

A data-driven analysis of energy balance closure across FLUXNET research sites: The role of landscape scale heterogeneity

The energy balance at most surface-atmosphere flux research sites remains unclosed. The mechanisms underlying the discrepancy between measured energy inputs and outputs across the global FLUXNET tower network are still under debate. Recent reviews have identified exchange processes and turbulent motions at large spatial and temporal scales in heterogeneous landscapes as the primary cause of the lack of energy balance closure at some intensively-researched sites, while unmeasured storage terms cannot be ruled out as a dominant contributor to the lack of energy balance closure at many other sites. We analyzed energy balance closure across 173 ecosystems in the FLUXNET database and explored the relationship between energy balance closure and landscape heterogeneity using MODIS products and GLOBEstat elevation data. Energy balance closure per research site (CEB,s) averaged 0.84±0.20, with best average closures in evergreen broadleaf forests and savannas (0.91–0.94) and worst average closures in crops, deciduous broadleaf forests, mixed forests and wetlands (0.70–0.78). Half-hourly or hourly energy balance closure on a percent basis increased with friction velocity (u*) and was highest on average under near-neutral atmospheric conditions. CEB,s was significantly related to mean precipitation, gross primary productivity and landscape-level enhanced vegetation index (EVI) from MODIS, and the variability in elevation, MODIS plant functional type, and MODIS EVI. A linear model including landscape-level variability in both EVI and elevation, mean precipitation, and an interaction term between EVI variability and precipitation had the lowest Akaike's information criterion value. CEB,s in landscapes with uniform plant functional type approached 0.9 and CEB,s in landscapes with uniform EVI approached 1. These results suggest that landscape-level heterogeneity in vegetation and topography cannot be ignored as a contributor to incomplete energy balance closure at the flux network level, although net radiation measurements, biological energy assimilation, unmeasured storage terms, and the importance of good practice including site selection when making flux measurements should not be discounted. Our results suggest that future research should focus on the quantitative mechanistic relationships between energy balance closure and landscape-scale heterogeneity, and the consequences of mesoscale circulations for surface-atmosphere exchange measurements.

Modelling carbon storage in highly fragmented and human-dominated landscapes: Linking land-cover patterns and ecosystem models

To extend coupled human–environment systems research and include the ecological effects of land-use and land-cover change and policy scenarios, we present an analysis of the effects of forest patch size and shape and landscape pattern on carbon storage estimated by BIOME-BGC. We evaluate the effects of including within-patch and landscape-scale heterogeneity in air temperature on carbon estimates using two modelling experiments. In the first, we combine fieldwork, spatial analysis, and BIOME-BGC at a 15-m resolution to estimate carbon storage in the highly fragmented and human-dominated landscape of Southeastern Michigan, USA. In the second, we perform the same analysis on 12 hypothetical landscapes that differ only in their degree of fragmentation. For each experiment we conduct four air-temperature treatments, three guided by field-based data and one empirically informed by local National Weather Service station data. The three field data sets were measured (1) exterior to a forest patch, (2) from the patch edge inward to 60 m on east-, south-, and west-facing aspects, separately, and (3) interior to that forest patch. Our field-data analysis revealed a decrease in maximum air temperature from the forest patch edge to a depth of 80 m. Within-patch air-temperature values were significantly different ( α = 0.01) among transects (c.v. = 13.28) and for all measurement locations (c.v. = 30.58). Results from the first experiment showed that the interior treatment underestimated carbon storage by ∼8000 Mg C and the exterior treatment overestimated carbon storage by 30,000 Mg C within Dundee Township, Southeastern Michigan, when compared to a treatment that included within-patch heterogeneity. In the second experiment we found a logarithmic increase in carbon storage with increasing fragmentation ( r 2 = 0.91). While a number of other processes (e.g. altered disturbance frequency or severity) remain to be included in future experiments, this combined field and modelling study clearly demonstrated that the inclusion of within-patch and landscape heterogeneity, and landscape fragmentation, each have a strong effect on forest carbon cycling and storage as simulated by a widely used ecosystem process model.

Litter decomposition promotes differential feedbacks in an oligotrophic southern Everglades wetland

The differential accumulation or loss of carbon and nutrients during decomposition can promote differentiation of wetland ecosystems, and contribute to landscape-scale heterogeneity. Tree islands are important ecosystems because they increase ecological heterogeneity in the Everglades landscape and in many tropical landscapes. Only slight differences in elevation due to peat accumulation allow the differentiation of these systems from the adjacent marsh. Hydrologic restoration of the Everglades landscape is currently underway, and increased nutrient supply that could occur with reintroduction of freshwater flow may alter these differentiation processes. In this study, we established a landscape-scale, ecosystem-level experiment to examine litter decomposition responses to increased freshwater flow in nine tree islands and adjacent marsh sites in the southern Everglades. We utilized a standard litterbag technique to quantify changes in mass loss, decay rates, and phosphorus (P), nitrogen (N) and carbon (C) dynamics of a common litter type, cocoplum (Chrysobalanus icaco L.) leaf litter over 64 weeks. Average C. icaco leaf degradation rates in tree islands were among the lowest reported for wetland ecosystems (0.23 ± 0.03 yr−1). We found lower mass loss and decay rates but higher absolute mass C, N, and P in tree islands as compared to marsh ecosystems after 64 weeks. With increased freshwater flow, we found generally greater mass loss and significantly higher P concentrations in decomposing leaf litter of tree island and marsh sites. Overall, litter accumulated N and P when decomposing in tree islands, and released P when decomposing in the marsh. However, under conditions of increased freshwater flow, tree islands accumulated more P while the marsh accumulated P rather than mineralizing P. In tree islands, water level explained significant variation in P concentration and N:P molar ratio in leaf tissue. Absolute P mass increased strongly with total P load in tree islands (r 2 = 0.81). In the marsh, we found strong, positive relationships with flow rate. Simultaneous C and P accumulation in tree island and mineralization in adjacent marsh ecosystems via leaf litter decomposition promotes landscape differentiation in this oligotrophic Everglades wetland. However, results of this study suggest that variation in flow rates, water levels and TP loads can shift differential P accumulation and loss leading to unidirectional processes among heterogeneous wetland ecosystems. Under sustained high P loading that could occur with increased freshwater flow, tree islands may shift to litter mineralization, further degrading landscape heterogeneity in this system, and signaling an altered ecosystem state.