Distinct ecosystem types respond differentially to grazing exclosure

Here, we evaluate the ecosystem functioning and the ecosystems services supply of different vegetation types (grasslands, shrublands and woodlands) under contrasting management regimes by comparing a protected area with the surrounding landscape, which has been subjected to human disturbance in the Eastern Hills of Uruguay. We propose, based on functional attributes and vegetation physiognomy, a State and Transition Model for the dynamics of the grassland–woodland mosaic. We used remote sensing techniques to: (i) develop a land‐cover map of the study area based on supervised Landsat imagery classification, and (ii) compare attributes of the ecosystem functioning (productivity and seasonality) and service supply derived from the Normalized Difference Vegetation Index (NDVI) images provided by the moderate resolution imaging spectroradiometer (MODIS) sensor. The land‐cover map showed that grasslands and shrublands were the most extensive land covers in the study area. These vegetation types presented higher productivity, seasonality and ecosystem service supply, outside the protected area than inside it. On the other hand, woodlands showed higher productivity, ecosystem service supply and lower seasonality inside the protected area than outside of it. Two axes represented the grassland–woodland mosaic dynamic: (i) the mean annual and (ii) the intra‐annual coefficient of variation of the NDVI. Our results highlight that conservation of grasslands, shrublands and woodlands require different management strategies based on particular disturbance regimes like moderate grazing and controlled burns. Moderate disturbances may help to preserve ecosystem services provisioning in grasslands and shrublands. On the contrary, woodland conservation requires a more rigorous regime of protection against disturbances.     

Modeling landscape patterns of understory tree regeneration in the Pacific Northwest,USA

Abstract. Vegetation maps serve as the basis for spatial analysis of forest ecosystems and provide initial information for simulations of forest landscape change. Because of the limitations of current remote sensing technology, it is not possible to directly measure forest understory attributes across large spatial extents. Instead we used a predictive vegetation mapping approach to model Tsuga heterophylla and Picea sitchensis seedling patterns in a 3900‐ha landscape in the Oregon Coast Range, USA, as a function of Landsat TM imagery, aerial photographs, digital elevation models, and stream maps. Because the models explained only moderate amounts of variability (R² values of 0.24–0.56), we interpreted the predicted patterns as qualitative spatial trends rather than precise maps. P. sitchensis seedling patterns were tightly linked to the riparian network, with highest densities in coastal riparian areas. T. heterophylla seedlings exhibited complex patterns related to topography and overstory forest cover, and were also spatially clustered around patches of old‐growth forest. We hypothesize that the old growth served as refugia for this fire‐sensitive species following wildfires in the late 19th and early 20th centuries. Low levels of T. heterophylla regeneration in hardwood‐dominated forests suggest that these patches may succeed to shrublands rather than to conifer forest. Predictive models of seedling patterns could be developed for other landscapes where georeferenced inventory plots, remote sensing data, digital elevation models, and climate maps are available.     

The interplay between facilitation and habitat type drives spatial vegetation patterns in global drylands

The spatial configuration of vascular vegetation has been linked to variations in land degradation and ecosystem functioning in drylands. However, most studies on spatial patterns conducted to date have focused on a single or a few study sites within a particular region, specific vegetation types, or in landscapes characterized by a certain type of spatial patterns. Therefore, little is known on the general typology and distribution of plant spatial patterns in drylands worldwide, and on the relative importance of biotic and abiotic factors as predictors of their variations across geographical regions and habitat types. We analyzed 115 dryland plant communities from all continents except Antarctica to: 1) investigate the general typology of spatial patterns, and 2) assess the relative importance of biotic (plant cover, frequency of facilitation, soil amelioration, height of the dominant species) and abiotic (aridity, rainfall seasonality and sand content) factors as predictors of spatial patterns (median patch size, shape of patch‐size distribution and regularity) across contrasting habitat types (shrublands and grasslands). Precipitation during the warmest period and sand content were particularly strong predictors of plant spatial patterns in grasslands and shrublands, respectively. Facilitation associated with power‐law like and irregular spatial patterns in both shrublands and grasslands, although it was mediated by different mechanisms (respectively soil ammelioration and percentage of facilitated species). The importance of biotic attributes as predictors of the shape of patch‐size distributions declined with aridity in both habitats, leading to the emergence of more regular patterns under the most arid conditions. Our results expand our knowledge about patch formation in drylands and the habitat‐dependency of their drivers. They also highlight different ways in which facilitation affects ecosystem structure through the formation of plant spatial patterns.     

Synergistic influences of introduced herbivores and fire on vegetation change in northern Patagonia,Argentina

Question: We investigated how cattle and European hares, the two most widespread exotic herbivores in Patagonia, affect species composition, life‐form composition and community structure during the first 6 years of vegetation recovery following severe burning of fire‐resistant subalpine forests and fire‐prone tall shrublands. We asked how the effects of introduced herbivores on post‐fire plant community attributes affect flammability of the vegetation. Location: Nahuel Huapi National Park, northwest Patagonia, Argentina Methods: We installed fenced plots to exclude livestock and European hares from severely burned subalpine forests of Nothofagus pumilio and adjacent tall shrublands of N. antarctica. The former is an obligate seed reproducer, whereas the latter and all other woody dominants of the shrubland vigorously resprout after burning. Results: Repeated measures ANOVA of annual measurements over the 2001‐2006 period indicate that cattle and hare exclusion had significant but complex effects on the cover of graminoids, forbs, climber species and woody species in the two burned community types. Significant interactions between the effects of cattle and hares varied by plant life forms between the two communities, which implies that their synergistic effects are community dependent. Conclusions: Following severe fires, the combined effects of cattle and hares inhibit forest recovery and favour transition to shrublands dominated by resprouting woody species. This herbivore‐induced trend in vegetation structure is consistent with the hypothesis that the effects of exotic herbivores at recently burned sites contribute to an increase in the overall flammability of the Patagonian landscape.     

Quantifying mortality of tropical rain forest trees using high-spatial-resolution satellite data

Assessment of forest responses to climate change is severely hampered by the limited information on tree death on short temporal and broad spatial scales, particularly in tropical forests. We used 1‐m resolution panchromatic IKONOS and 0.7‐m resolution QuickBird satellite data, acquired in 2000 and 2002, respectively, to evaluate tree death rates at the La Selva Biological Station in old‐growth Tropical Wet Forest in Costa Rica, Central America. Using a calibration factor derived from ground inspection of tree deaths predicted from the images, we calculated a landscape‐scale annual exponential death rate of 2.8%. This corresponds closely to data for all canopy‐level trees in 18 forest inventory plots, each of 0.5 ha, for a mostly‐overlapping 2‐year period (2.8% per year). This study shows that high‐spatial‐resolution satellite data can now be used to measure old‐growth tropical rain forest tree death rates, suggesting many new avenues for tropical forest ecology and global change research.     

Measuring canopy loss and climatic thresholds from an extreme drought along a fivefold precipitation gradient across Texas

Globally, trees are increasingly dying from extreme drought, a trend that is expected to increase with climate change. Loss of trees has significant ecological, biophysical, and biogeochemical consequences. In 2011, a record drought caused widespread tree mortality in Texas. Using remotely sensed imagery, we quantified canopy loss during and after the drought across the state at 30‐m spatial resolution, from the eastern pine/hardwood forests to the western shrublands, a region that includes the boundaries of many species ranges. Canopy loss observations in ~200 multitemporal fine‐scale orthophotos (1‐m) were used to train coarser Landsat imagery (30‐m) to create 30‐m binary statewide canopy loss maps. We found that canopy loss occurred across all major ecoregions of Texas, with an average loss of 9.5%. The drought had the highest impact in post oak woodlands, pinyon‐juniper shrublands and Ashe juniper woodlands. Focusing on a 100‐km by ~1,000‐km transect spanning the State's fivefold east–west precipitation gradient (~1,500 to ~300 mm), we compared spatially explicit 2011 climatic anomalies to our canopy loss maps. Much of the canopy loss occurred in areas that passed specific climatic thresholds: warm season anomalies in mean temperature (+1.6°C) and vapor pressure deficit (VPD, +0.66 kPa), annual percent deviation in precipitation (?38%), and 2011 difference between precipitation and potential evapotranspiration (?1,206 mm). Although similarly low precipitation occurred during the landmark 1950s drought, the VPD and temperature anomalies observed in 2011 were even greater. Furthermore, future climate data under the representative concentration pathway 8.5 trajectory project that average values will surpass the 2011 VPD anomaly during the 2070–2099 period and the temperature anomaly during the 2040–2099 period. Identifying vulnerable ecological systems to drought stress and climate thresholds associated with canopy loss will aid in predicting how forests will respond to a changing climate and how ecological landscapes will change in the near term.     

Spatial early warning signals for impending regime shifts: A practical framework for application in real‐world landscapes

Prediction of ecosystem response to global environmental change is a pressing scientific challenge of major societal relevance. Many ecosystems display nonlinear responses to environmental change, and may even undergo practically irreversible ‘regime shifts’ that initiate ecosystem collapse. Recently, early warning signals based on spatiotemporal metrics have been proposed for the identification of impending regime shifts. The rapidly increasing availability of remotely sensed data provides excellent opportunities to apply such model‐based spatial early warning signals in the real world, to assess ecosystem resilience and identify impending regime shifts induced by global change. Such information would allow land‐managers and policy makers to interfere and avoid catastrophic shifts, but also to induce regime shifts that move ecosystems to a desired state. Here, we show that the application of spatial early warning signals in real‐world landscapes presents unique and unexpected challenges, and may result in misleading conclusions when employed without careful consideration of the spatial data and processes at hand. We identify key practical and theoretical issues and provide guidelines for applying spatial early warning signals in heterogeneous, real‐world landscapes based on literature review and examples from real‐world data. Major identified issues include (1) spatial heterogeneity in real‐world landscapes may enhance reversibility of regime shifts and boost landscape‐level resilience to environmental change (2) ecosystem states are often difficult to define, while these definitions have great impact on spatial early warning signals and (3) spatial environmental variability and socio‐economic factors may affect spatial patterns, spatial early warning signals and associated regime shift predictions. We propose a novel framework, shifting from an ecosystem perspective towards a landscape approach. The framework can be used to identify conditions under which resilience assessment with spatial remotely sensed data may be successful, to support well‐informed application of spatial early warning signals, and to improve predictions of ecosystem responses to global environmental change.     

Testing the Wiegand–Milton model: A long‐term experiment to understand mechanisms driving vegetation dynamics in arid shrublands

The Wiegand and Milton (1996) simulation model predicts that vegetation dynamics in arid shrublands are characterized by event‐driven stochasticity (weather events), and demographic inertia (persistence of a species in a community) that lead to a lagged response in vegetation compositional change. Slow plant growth is one of the mechanisms driving slow vegetation change. We test this model at the same location (Tierberg Long‐term Ecological Research site) on which the model was based. Three dwarf shrub species, differing in palatability, were tracked over 25 years (1988–2014) at two levels of the past herbivory (pre‐1960) and three levels of the present herbivory (post‐1988). In the period between 1960 and 1988, all sites were grazed at the recommended agricultural stocking rate. For each species, plant density and a number of size attributes (basal diameter, height, canopy area) were surveyed. Analyses using a two‐way Analysis of Covariance (ANCOVA) took initial starting size into consideration. As the model predicted, event‐driven stochasticity (rainfall) resulted in an increase in density of the smaller size classes following a single large recruitment event across all grazing regimes for the palatable and unpalatable species. Size‐class distribution curve types remained unchanged illustrating that population demography remains unaffected for long periods and responses are slow (lagged response). Slow plant growth was evident in that there were no changes in height, canopy area, or density under present grazing regimes over the 25‐year period. Palatable species had a reduced canopy area and density compared to unpalatable species. Our findings provide empirical evidence supporting the predictions of the Wiegand and Milton (1996) model, notably event‐driven stochasticity, demographic inertia, and a lagged response in vegetation change in arid shrublands. In addition, our results support the model assumption of the significance of slow growth in long‐lived plant species and the influence of grazing regime.     

Regional scale relationships between ecosystem structure and functioning: the case of the Patagonian steppes

Aims 1. To characterize ecosystem functioning by focusing on above‐ground net primary production (ANPP), and 2. to relate the spatial heterogeneity of both functional and structural attributes of vegetation to environmental factors and landscape structure. We discuss the relationship between vegetation structure and functioning found in Patagonia in terms of the capabilities of remote sensing techniques to monitor and assess desertification. Location Western portion of the Patagonian steppes in Argentina (39°30′ S to 45°27′ S). Methods We used remotely‐sensed data from Landsat TM and AVHRR/NOAA sensors to characterize vegetation structure (physiognomic units) and ecosystem functioning (ANPP and its seasonal and interannual variation). We combined the satellite information with floristic relevés and field estimates of ANPP. We built an empirical relationship between the Landsat TM‐derived normalized difference vegetation index (NDVI) and field ANPP. Using stepwise regressions we explored the relationship between ANPP and both environmental variables (precipitation and temperature surrogates) and structural attributes of the landscape (proportion and diversity of different physiognomic classes (PCs)). Results PCs were quite heterogeneous in floristic terms, probably reflecting degradation processes. Regional estimates of ANPP showed differences of one order of magnitude among physiognomic classes. Fifty percent of the spatial variance in ANPP was accounted for by longitude, reflecting the dependency of ANPP on precipitation. The proportion of prairies and semideserts, latitude and, to a lesser extent, the number of PCs within an 8 × 8 km cell accounted for an additional 33% of the ANPP variability. ANPP spatial heterogeneity (calculated from Landsat TM data) within an 8 × 8 km cell was positively associated with the mean AVHRR/NOAA NDVI and with the diversity of physiognomic classes. Main conclusions Our results suggest that the spatial and temporal patterns of ecosystem functioning described from ANPP result not only from water availability and thermal conditions but also from landscape structure (proportion and diversity of different PCs). The structural classification performed using remotely‐sensed data captured the spatial variability in physiognomy. Such capability will allow the use of spectral classifications to monitor desertification.     

Environmental drivers of femaleness of an inter‐Andean monoecious shrub

Hetero‐and conspecific interactions, nutrient availability, climate, habitat heterogeneity, and disturbances can generate variation and spatial patterns of femaleness in plants. We assessed whether year, site, plant size, plant density, and canopy area of conspecific neighbors influenced the expression and spatial aggregation of femaleness in Croton aff. wagneri, a monoecious shrub from dry shrublands of the inter‐Andean valleys in Ecuador. We georeferenced in two sites (1,700 and 1,400 m.a.s.l) in five 10 × 10 m plots, within each site, the position of each Croton reproductive plant during first part of flowering season in two years, and measured their height, length, and width. The femaleness index of each plant was determined by the number of female and male buds and flowers. Plant density was determined for each plot, along with the number of neighbors and the summed canopy area of conspecific neighbors (at 1.0, 2.0, and 2.5 m radius, and the five closest plants) from each focal plant. Croton´s femaleness at the lower elevation site was greater than at the higher elevation site and increased with plant size and with canopy of the closest five neighbors. Soil at the lower elevation site had higher temperatures and lower water content. Aggregate patterns of femaleness were found in more plots at the lower elevation site. Our results indicate that location, plant size, and canopies of conspecific neighbors of Croton can affect femaleness and its aggregation and support the hypothesis that femaleness can be influenced by facilitative interactions. Abstract in Spanish is available with online material.     

Multiple periglacial refugia in the Patagonian steppe and post‐glacial colonization of the Andes: the phylogeography of Calceolaria polyrhiza

Aim We perform a phylogeographical study of an endemic Patagonian herbaceous plant to assess whether geographical patterns of genetic variation correspond to in situ Pleistocene survival or to glacial retreat and post‐glacial expansion. We also seek to determine the locations of potential glacial refugia and post‐glacial colonization routes. Location Southern Andes and Patagonian steppe. Methods We used Calceolaria polyrhiza, a widely distributed Patagonian herbaceous plant that occurs mainly in the understorey of Nothofagus rain forests and in the arid Patagonian steppe, as our model system. The chloroplast intergenic spacer trnH–psbA was sequenced for 590 individuals from 68 populations. Sequence data were analysed using phylogenetic (maximum parsimony, maximum likelihood and Bayesian inference) and population genetic (spatial analyses of molecular variance, mismatch distributions and neutrality tests) methods. Nested clade phylogeographic analyses, and divergence time estimates using a calibrated molecular clock, were also conducted. Results A total of 27 haplotypes identified in the present study clustered into four primary genealogical lineages, revealing three significant latitudinal phylogeographical breaks. The two high Andean lineages probably split first, during the late Miocene, and the Patagonian lineage split around 4 Ma, coincident with the establishment of the Patagonian steppe. Within each haplogroup, major diversification occurred in the Pleistocene. The Patagonian groups show a pattern consistent with a rapid post‐glacial expansion and colonization of the Andean flanks, achieved independently by four lineages. The highest haplotype diversity was found along a longitudinal transect that is remarkably congruent with the limit of the ice‐sheet extension during the Greatest Patagonian Glaciation. A north‐east expansion is evident, which is probably associated with the ‘Arid Diagonal’ fluctuations. Main conclusions Glacial climate fluctuations had a substantial impact on the diversification, distribution and demography of the study species. A scenario of multiple periglacial Pleistocene refugia and subsequent multiple recolonization routes, from eastern Patagonia to the Andean flanks, may explain the phylogeographical patterns observed. However, current genetic structure also preserves the imprints of older events that probably occurred in the Miocene and Pliocene, providing evidence that multiple processes, operating at different spatial and temporal scales, have moulded biodiversity in Patagonia.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Have we been successful? Monitoring horizontal forest complexity for forest restoration projects

Forest management today often seeks to restore ecological integrity and enhance human well‐being by increasing forest complexity, resilience, and functionality. However, effective and financially expedient monitoring of forest complexity is challenging. In this study, we developed a practical and inexpensive technique to measure horizontal forest complexity. This monitoring method uses intuitively understandable data (imagery) and facilitates stakeholder participation in the adaptive management process within collaborative projects. We used this technique to determine if current restoration projects are successfully achieving their spatial restoration goals. We focused on the Colorado Front Range Landscape Restoration Initiative (CFRLRI) as a representative of the typical collaborative restoration projects underway in formerly fire‐dependent dry conifer forests. The developed monitoring method is practical and cost‐effective by using free aerial imagery to map, quantify, and analyze the distribution of canopy cover pre‐ and post‐treatment. We found the CFRLRI has successfully reduced canopy cover (from 44 to 26% on average) and increased some aspects of horizontal forest complexity. The application of these monitoring techniques has allowed the CFRLRI collaborative group to objectively quantify changes to horizontal forest complexity, and has facilitated stakeholder communication about forest spatial patterns. These methods could be adapted for use by other similar forest restoration projects around the world by utilizing increasingly available satellite or aerial imagery.     

Forest bird diversity in Mediterranean areas affected by wildfires: a multi-scale approach

Fire is a key mechanism creating and maintaining habitat heterogeneity in Mediterranean landscapes by turning continuous woody landscapes into mosaics of forests and shrublands. Due to the long historical role of fires in the Mediterranean, we hypothesised a moderate negative effect of this type of perturbation on forest bird distribution at a landscape level. We conducted point bird censuses in Aleppo pine forest patches surrounded by burnt shrublands and studied the relationships between three ecological groups of bird species (forest canopy species, forest understorey species, and ubiquitous species) and the features of local habitat, whole patch and surrounding landscape. We used a multi-scale approach to assess the effects of landscape variables at increasing spatial scales on point bird richness. Regarding local habitat components, canopy species were positively associated with tall pines while understorey species with the cover of shrubs and plants from holm-oak forests. Forest birds were positively related to patch size and irregular forest shapes, that is, with high perimeter/size ratios. Thus, these species did not seem to perceive edges as low quality but rather favourable microhabitats. We did not detect any negative effect of isolation or cover of woodlands in the landscape on the presence of forest species after local habitat factors had been accounted for. Finally, only local habitat factors entered the model for ubiquitous species. We suggest that mosaic-like landscapes shaped by fires in the Mediterranean basin are not strongly associated with negative effects fragmentation on forest birds other than those related with habitat loss.     

Directional climate change and potential reversal of desertification in arid and semiarid ecosystems

Our objective was to determine if long‐term increases in precipitation can maintain grasslands susceptible to desertification, and initiate a reversal of historic regime shifts on desertified shrublands. Perennial grass production and species richness in a multi‐year wet period were hypothesized to be greater than expected based on precipitation in a sequence of dry years. These responses were expected to differ for grasslands and shrublands with different dominant species and topo‐edaphic properties. Long‐term trends in desertification were documented using vegetation maps beginning in 1858, 1915, 1928, and 1998). These trends were compared with herbaceous and woody species responses to a sequence of dry (1994–2003) and wet years (2004–2008) for two grassland (uplands, playas) and three desertified shrubland types (honey mesquite, creosotebush, tarbush) in the Chihuahuan Desert. Analyses showed that both types of grasslands decreased in spatial extent since 1858 whereas areas dominated by mesquite or creosotebush increased. Production of upland grasslands in the wet period was greater than expected based on responses during the dry period whereas the relationships between species richness and precipitation was the same for both periods. Precipitation was not important to responses in playa grasslands in either period. For all ecosystem types, the production response in wet years primarily was an increase in herbaceous plants, and the most pronounced responses occurred on sandy sites (upland grasslands, mesquite shrubland). Results suggest that multiple wet years are needed to initiate a sequence of grass establishment and survival processes that can maintain upland grasslands without management inputs and lead to a state change reversal in desertified shrublands. Restoration strategies need to take advantage of opportunities provided by future climates while recognizing the importance of ecosystem type.     

French broom (Teline monspessulana) invasion in south-central Chile depends on factors operating at different spatial scales

The interaction between plant attributes, environmental factors and the history of human intervention determines which species show a more invasive behavior and which areas are more likely to be invaded. Identifying which factors favor the invasion and which constraint it, remains an essential goal for understanding the process of invasion and to provide a stronger scientific basis for designing management actions to reduce susceptibility to invasion. In this paper we analyze the relative role of different environmental factors in plant invasion, in particular the comparison between human versus climatic and biotic variables at different spatial scales. Furthermore, we show how these factors interact to influence the distribution and abundance of an invasive plant. We have built mixed models to explain the presence and abundance of a renowned invasive species in the south-central area of Chile, Teline monspessulana (French broom, Fabaceae). We recorded the presence and cover of this invasive shrub in a series of transects along the main roads of the study area. For each transect we recorded variables at different spatial scales related to climate and land use. We found that the presence and cover of T. monspessulana are strongly related to proximity to roads, urban areas and locations with higher rainfall. Although this shrub is present in most of the landscape, it is most abundant in shrublands and forest plantations. In a complex human-dominated landscape matrix, shrublands and forest plantations act as a source of propagules from which new areas can be invaded. Interestingly, the presence of this species can be better predicted by combining predictors taken at two hierarchical levels, the so-called transect and plot scales, whereas the species abundance only depends on factors at the plot level. The presence and abundance of this species is highly dependent on human disturbance, and the occurrence of certain landscape elements closely linked to land management is critical to the performance of this species. Maintaining a dense tree canopy cover and reducing the use of fire could help reduce the rate of expansion of this species across the landscape.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

Landscape responses to a century of land use along the northern Patagonian forest-steppe transition

Land use history reconstructions in temperate regions of the Northern Hemisphere indicate that periods of deforestation are often followed by natural afforestation, so that the long-term outcome at the landscape level will be a balance of retractions and advances of plant communities associated with varying local land uses. During the last decades of the XIX century, large forest areas were cleared in Northwestern Patagonia to open farmland. In this article, we compared historical land use/land cover maps with land cover maps derived from Landsat images to analyze the factors that may have influenced the dynamics of land cover change of the forest-steppe ecotone during the last 100 years. Our results indicate that Patagonian forests underwent a rapid initial recovery after the extensive fires of last century, replacing mainly shrublands. More than 50% of the old burns are currently covered by forests, and modern fires affect areas characterized by fire-prone vegetation. Whereas natural afforestation is an ongoing process positively associated with moisture, the rate of forest losses has increased during the last three decades, concentrating on xeric aspects and the vicinity of roads. We conclude that the outcome of the dynamics between fire-intolerant forests and fire-prone plant communities will largely depend on human-related activities, modeled by structural features of the landscape (i.e., topography, dominant winds), and processes triggered by past land uses.     

Heteroskedasticity as a leading indicator of desertification in spatially explicit data

Regime shifts are abrupt transitions between alternate ecosystem states including desertification in arid regions due to drought or overgrazing. Regime shifts may be preceded by statistical anomalies such as increased autocorrelation, indicating declining resilience and warning of an impending shift. Tests for conditional heteroskedasticity, a type of clustered variance, have proven powerful leading indicators for regime shifts in time series data, but an analogous indicator for spatial data has not been evaluated. A spatial analog for conditional heteroskedasticity might be especially useful in arid environments where spatial interactions are critical in structuring ecosystem pattern and process. We tested the efficacy of a test for spatial heteroskedasticity as a leading indicator of regime shifts with simulated data from spatially extended vegetation models with regular and scale‐free patterning. These models simulate shifts from extensive vegetative cover to bare, desert‐like conditions. The magnitude of spatial heteroskedasticity increased consistently as the modeled systems approached a regime shift from vegetated to desert state. Relative spatial autocorrelation, spatial heteroskedasticity increased earlier and more consistently. We conclude that tests for spatial heteroskedasticity can contribute to the growing toolbox of early warning indicators for regime shifts analyzed with spatially explicit data.