Old-growth forest landscape transitions from pre-European settlement to present

We conducted a multi-temporal spatial analysis of forest cover for a 9600 ha landscape in northern Wisconsin, U.S.A., using data from pre-European settlement (1860s), post-settlement (1931), and current (1989) periods. Using GIS we have shown forest landscape changes and trajectories that have been generally described in aggregate for the norther Great Lake States region. We created the pre-European settlement map from the witness tree data of the original federal General Land Office survey notes. The 1931 cover was produced from the Wisconsin Land Economic Inventory, and the 1989 cover map was based on color infrared photography. We used GIS to analyze 1) land area occupied by different forest types at different dates, 2) temporal transitions between dates and their driving proceses, and 3) successional trajectories with landforms and spatial associations of forest types. Over the 120 year period, forest cover has changed from a landscape dominated by old-growth hemlock (Tsuga canadensis) and hardwood forests (Acer saccharum, Betula alleghaniensis) to largely second-growth hardwoods and conifers. The former dominant hemlock is largely eliminated from the landscape. From 1860 to 1931, large-scale disturbances associated with logging were the dominant processes on the landscape. Early successional forest types covered much of the landscape by the 1930s. From 1931 to 1989, succession was the dominant process driving forest transitions as forest types succeeded to a diverse group of upland hardwood and conifer forest types. If successional trajectories continue, a more homogeneous landscape may develop comprised of both a northern hardwood type dominated by sugar maple, and a boreal conifer/hardwood forest.     

Exploring Patterns of Exotic Earthworm Distribution in a Temperate Hardwood Forest in South-Central New York, USA

Exotic earthworms invading forests in Canada and northeastern United States that were naturally devoid of large detritivores cause major changes in ecosystem function. To assess their long-term impacts, studies are needed to elucidate the factors that control the patterns of earthworm invasion at the landscape level. We analyzed the distribution patterns of exotic earthworms in a northern hardwood forest in south-central New York (USA), as explained by landscape variables thought to be important in determining earthworm distribution. Forest type, slope angle, elevation, and the distance to agricultural clearings and wet refugia were significant predictors of earthworm presence, whereas local wetness index and the distance to streams and roads were not. Forest type and distance to agricultural clearings were the two most significant predictors. Our data suggest that areas close to agricultural clearings, dominated by mixed hardwoods, and located towards valley bottoms or on gentle slopes are very likely to support communities of exotic earthworms. Steeper slopes, areas dominated by American beech or eastern hemlock, and locations in the core of extensive forest landscapes have lower probabilities of invasion by exotic earthworms. When applied to a nearby area, our statistical model correctly predicted earthworm presence for 67% of 377 sampling points. Most of the mistakes were incorrect predictions of earthworm absence, suggesting that our statistical model slightly underestimated earthworm presence, possibly because of the pervasive influence of active agricultural fields adjacent to the test site.     

Mixed-severity fire regime in a high-elevation forest of Grand Canyon,Arizona, USA

Fire regime characteristics of high-elevation forests on the North Rim of the Grand Canyon, Arizona, were reconstructed from fire scar analysis, remote sensing, tree age, and forest structure measurements, a first attempt at detailed reconstruction of the transition from surface to stand-replacing fire patterns in the Southwest. Tree densities and fire-/non-fire-initiated groups were highly mixed over the landscape, so distinct fire-created stands could not be delineated from satellite imagery or the oldest available aerial photos. Surface fires were common from 1700 to 1879 in the 4,400 ha site, especially on S and W aspects. Fire dates frequently coincided with fire dates measured at study sites at lower elevation, suggesting that pre-1880 fire sizes may have been very large. Large fires, those scarring 25% or more of the sample trees, were relatively infrequent, averaging 31 years between burns. Four of the five major regional fire years occurred in the 1700s, followed by a 94-year gap until 1879. Fires typically occurred in significantly dry years (Palmer Drought Stress Index), with severe drought in major regional fire years. Currently the forest is predominantly spruce-fir, mixed conifer, and aspen. In contrast, dendroecological reconstruction of past forest structure showed that the forest in 1880 was very open, corresponding closely with historical (1910) accounts of severe fires leaving partially denuded landscapes. Age structure and species composition were used to classify sampling points into fire-initiated and non-fire-initiated groups. Tree groups on nearly 60% of the plots were fire-initiated; the oldest such groups appeared to have originated after severe fires in 1782 or 1785. In 1880, all fire-initiated groups were less than 100 years old and nearly 25% of the groups were less than 20 years old. Non-fire-initiated groups were significantly older (oldest 262 years in 1880), dominated by ponderosa pine, Douglas-fir, or white fir, and occurred preferentially on S and W slopes. The mixed-severity fire regime, transitioning from lower-elevation surface fires to mixed surface and stand-replacing fire at higher elevations, appeared not to have been stable over the temporal and spatial scales of this study. Information about historical fire regime and forest structure is valuable for managers but the information is probably less specific and stable for high-elevation forests than for low-elevation ponderosa pine forests.This revised version was published online in May 2005 with corrections to the Cover Date.     

Post-storm surveys reveal large-scale spatial patterns and influences of site factors,forest structure and diversity in endemic bark-beetle populations

The storm that struck France on december 26^th and 28^th 1999 felled 140 million m³ of timber and had a high economic, social and landscape impact. This event offered the opportunity to study large-scale patterns in populations of forest insect pests that would benefit from the abundant breeding material. A large-scale survey was carried out in France in 2000 to sample the most frequently observed species developing on spruce (Ips typographus, Pityogene schalcographus) and pine (Tomicus piniperda, Ips sexdentatus) in 898 locations distributed throughout wind-damaged areas. The local abundance of each species scored on a 0 to 5 scale was analysed using geostatistical estimators to explore the extent and intensity of spatial autocorrelation, and was related to site, stand, and neighbourhood landscape metrics of the forest cover (in particular the interconnection with broadleaf forest patches) found within dispersal distance. All species but I. sexdentatus, which was much less abundant, displayed large-scale spatial dependence and regional variations in abundance. Lower infestation levels per tree (windfalls and standing trees) were observed in stands with a high proportion of wind-damaged trees, which was interpreted as the result of beetles distributing themselves among the available breeding material. More infestations were observed in wind-broken trees as compared to wind-felled trees. More importantly, populations showed significant relationships with the structure of coniferous stands (in particular with the number of coniferous patches). T. piniperda population levels were negatively correlated to the amount of coniferous edge shared with broadleaf forest patches, possibly because of the disruptive effect of non-host volatiles on host-finding processes at the landscape-scale. The differences observed between species regarding patterns and relationships to site, stand, and forest cover characteristics are discussed in relation to the ecological characteristics of each species.     

Predicting Southern Appalachian overstory vegetation with digital terrain data

Vegetation in mountainous regions responds to small-scale variation in terrain, largely due to effects on both temperature and soil moisture. However there are few studies of quantitative, terrain-based methods for predicting vegetation composition. This study investigated relationships between forest composition, elevation, and a derived index of terrain shape, and evaluates methods for predicting forest composition. Trees were measured on 406 permanent plots within the boundaries of the Coweeta Hydrologic Lab, located in the Southern Appalachian Mountains of western North Carolina, USA. All plots were in control watersheds, without human or major natural disturbance since 1923. Plots were 0.08 ha and arrayed on transects, with approximately 380 meters between parallel transects. Breast-height diameters were measured on all trees. Elevation and terrain shape (cove, ridge, sideslope) were estimated for each plot. Density (trees/ha) and basal area were summarized by species and by forest type (cove, xeric oak-pine, northern hardwoods, and mixed deciduous). Plot data were combined with a digital elevation data (DEM), and a derived index of terrain shape at two sampling resolutions: 30 m (US Geological Survey), and 80 m (Defense Mapping Agency) sources. Vegetation maps were produced using each of four different methods: 1) linear regression with and without log transformations against elevation and terrain variables combined with cartographic overlay, 2) kriging, 3) co-kriging, and 4) a mosaic diagram. Predicted vegetation was compared to known vegetation at each of 77 independent, withheld data points, and an error matrix was determined for each mapping method.We observed strong relationships between some species and elevation and/or terrain shape. Cove and xeric oak/pine species basal areas were positively and negatively related to concave landscape locations, respectively, while species typically found in the mixed deciduous and northern hardwood types were not. Most northern hardwood species occurred more frequently and at higher basal areas as elevation increased, while most other species did not respond to elevation. The regression and mosaic diagram mapping approaches had significantly higher mapping accuracies than kriging and co-kriging. There were significant effects of DEM resolution on map accuracy, with maps based on 30 m DEM data significantly more accurate than those based on 80 m data. Taken together, these results indicate that both the mapping method and terrain data resolution significantly affect the resultant vegetation maps, even when using relatively high resolution data. Landscape or regional models based on 100 m or lower resolution terrain data may significantly under-represent terrain-related variation in vegetation composition.     

Disturbance and landscape dynamics in the Chequamegon National Forest Wisconsin,USA, from 1972 to 2001

Land uses, especially harvesting and road building, are considered to be the primary cause of forest fragmentation in many parts of the world. To test this perception, we (1) quantified changes and rates of change in vegetative composition and structure within the Washburn Ranger District in northern Wisconsin using Landsat images, (2) examined changes in landscape structure, (3) assessed changes within the area of road influence (ARI), and (4) investigated changes in landscape composition and structure within the context of forest management activities. Our landscape classifications included six dominant cover types: mixed hardwood (MH), jack pine (JP), red pine (RP), mixed hardwood/conifer (MHC), non-forested bare ground (NFBG), and regenerating forest or shrub (RFS). Increases in NFBG and RFS, by 196% and 28% respectively, reflect expansion of the pine-barrens. Windthrow in the mature hardwoods during the late 1970s and jack pine budworm outbreaks during the mid-1990s correlated with decreases in those classes over the corresponding intervals. A 69% decrease in mean patch size and a 60% increase in edge density reflect increased fragmentation. An inverse relationship existed between the compositional trends of forested (excluding JP) cover types and RFS and NFBG cover types. ARI covered 8% of the landscape affecting species composition within the MH, RFS, and NFBG. Results from this study are key in assessing the links between management activities and ecological consequences and thereby facilitate adaptive management.This revised version was published online in May 2005 with corrections to the Cover Date.     

Response of North American ecosystem models to multi-annual periodicities in temperature and precipitation

Ecosystem models typically use input temperature and precipitation data generated stochastically from weather station means and variances. Although the weather station data are based on measurements taken over a few decades, model simulations are usually on the order of centuries. Consequently, observed periodicities in temperature and precipitation at the continental scale that have been correlated with largscale forcings, such as ocean-atmosphere dynamics and lunar and sunspot cycles, are ignored. We investigated how these natural climatic fluctuations affect aboveground biomass in ecosystem models by incorporating some of the more pronounced continental-scale cycles in temperature (4, 11, 80, 180 year periods) and precipitation (11 and 19 year periods) into models of three North American forests (using LINKAGES) and one North American grassland (using STEPPE). Even without inclusion of periodicities in climate, long-term dynamics of these models were characterized by internal frequencies resulting from vegetation birth, growth and death processes. Our results indicate that long-term temperature cycles result in significantly lower predictions of forest biomass than observed in the control case for a forest on a biome transition (northern hardwoods/boreal forest). Lower-frequency, higher-amplitude temperature oscillation caused amplification of forest biomass response in forests containing hardwood species. Shortgrass prairie and boreal ecosystems, dominated by species with broad stress tolerance ranges, were relatively insensitive to climatic oscillations. Our results suggest periodicities in climate should be incorporated within long-term simulations of ecosystems with strong internal frequencies, particularly for systems on biome transitions.     

Advance regeneration and seed banking of woody plants in Ohio pine plantations: Implications for landscape change

Silviculturally-managed pine plantations within southern Ohio are chronically disturbed patches of introduced vegetation distinct from the surrounding matrix of hardwood forest. To determine the successional pathways by which such pine stands might blend back into the hardwood forest matrix under different types of silvicultural management, we determined the current status of hardwood regeneration under 24 pine stands. Stands of Pinus virginiana (Virginia pine) had the highest density of hardwood seedlings and saplings (20,560 stems ha^-1) whereas P. strobus (white pine) stands averaged only 7090 hardwood stems ha^-1; P. resinosa (red pine) stands were intermediate. The most abundant hardwood seedling and sapling species under pine canopies were Acer rubrum and Cornus florida. DCA ordination of the seedling + sapling assemblages clustered most of the P. resinosa and P. strobus stands in the center of the ordination along with a group of species which are common in second-growth forests of the area. P. virginiana stands, in contrast, were scattered throughout the ordination space. Most of the woody species common in second-growth forests of the region were also common in the pine understory. Multiple regression indicated that large plantations with deeper litter, higher soil pH and lower total hardwood density had the greatest abundance of mesic-site species in the understory. This relationship did not hold for P. resinosa stands, however, due to more frequent and intense silvicultural intervention. The seed bank was not an important source of woody seedlings to the understory assemblage under intact pine plantations. The vegetation of 1–4 yr old clear-cut sites was dominated by wind and bird dispersed species which were generally absent from the understory of intact plantations. We conclude these chronically disturbed planted patches will revert to matrix vegetation faster if the disturbance is allowed to end in a gradual manner through stand senescence than if it is abruptly ended by clear-cutting.     

Presettlement landscape heterogeneity: Evaluating grain of resolution using General Land Office Survey data

General Land Office Survey (GLOS) records from the A.D. 1840s provide data for quantitative characterization of presettlement vegetation across western Mackinac County, Michigan, located within the mixed conifer-northern hardwoods forest region. We analyzed data from land survey plat maps and 1958 bearing, witness, and line trees from 162 surveyed section and quarter-section corners in order to map vegatation cover types at a level of spatial resolution appropriate for characterizing landscape heterogeneity using standard landscape ecological metrics. As also demonstrated by a number of both classic and contemporary plant-ecological studies, the distribution of landforms, soils properties, hydrology, and location of fire breaks all contribute to the heterogeneity in vegetation observed at a landscape scale in the region. Through a series of spatial landscape analyses with differing grain of resolution, in this study we determine that a grid cell size of 65 ha (0.5 mi×0.5 mi or 0.25 mi²) to 259 ha (1 mi²) gives a conservative characterization of landscape heterogeneity using standard metrics and is therefore appropriate for use of GLOS data to study historical landscape changes.     

Aspen persistence near the National Elk Refuge and Gros Ventre Valley elk feedgrounds of Wyoming, USA

We investigated aspen (Populus tremuloides)regeneration in the Gros Ventre River Valley, the National Elk Refuge and a small part of Grand Teton National Park, Wyoming, USA to see if elk (Cervus elaphus) browsing was as damaging as previously thought. We conducted a landscape-scale survey to assess aspen regeneration across gradients of wintering elk concentrations using 68 randomly selected aspen stands in the 1090 km² study area. Forty-four percent of the stands sampled supported some newer regeneration that had reached the canopy. There were no significant differences of regeneration across elk winter range classification (p=0.25) or distance from feedgrounds (p=0.96). However, a multiple linear regression found that the concentration of elk was one of several important predictors of successful aspen regeneration (p=0.005, R ²=0.36). Our results suggest that stand-replacing regeneration occurs across the landscape at a variety of elk densities despite some trends of reduced regeneration under greater elk concentrations. We propose that high spatial and temporal variation and scattered patches of successful aspen regeneration characterize aspen persistence between periods of episodic regeneration and recruitment.     

Regeneration Strategies,Disturbance and Plant Interactions as Organizers of Vegetation Spatial Patterns in a Pine Forest

To determine how vegetation pattern in early successional forests may be related to plant traits and types of disturbance, we measured percent cover of individual taxa annually in a South Carolina Pinus elliottii forest, starting one year before, and ending four years after harvest and tree girdling disturbances were applied. The 17 most important taxa surveyed were grouped into four regeneration strategies chosen a priori, and the spatial patterns of these groups and of the soil were investigated using global variability, semivariograms and kriged maps. We also examined spatial correlations across years, across taxa, and between species and soil disturbance. Seed bank taxa represented by Dichanthelium spp. increased rapidly and formed large patches, and then quickly declined. Taxa that regenerate by newly dispersed seeds, represented by Rhus copallina and Rubus spp. occurred at first in a few patches, and became widespread later. Stump sprouters, represented by Quercus spp. and Myrica cerifera, had rapid increases in cover, but their spatial patterns were largely determined by their pre-disturbance patterns. Prunus serotina, which relies on both sprouting and dispersed seed, had moderate cover and a random distribution. Within-species temporal correlation of spatial pattern was lower in girdled than in harvested plots, and was not clearly related to regeneration strategy. Forest floor disturbance was patchy and affected the pattern of Dichanthelium spp. in the harvested plots. Negative correlations between herbs and woody plants in harvested plots reflected the role of biotic (i.e., successional) filters on vegetation pattern. Surprisingly, no spatial correlations were detected between the nitrogen fixer, Myrica cerifera and other taxa in this N-limited system. In comparing the spatial and temporal patterns, we found kriged maps more informative than analysis of semivariograms alone. The maps and correlation statistics demonstrated that regeneration traits, spatial patterns of soil disturbances, and interactions among taxa influence dynamics of the spatial patterns of the plants. We also demonstrated that disturbance types affected the importance and interactions among these three factors, and caused different spatial patterns of the plant taxa.     

Ecotonal changes and altered tree spatial patterns in lower mixed-conifer forests, Grand Canyon National Park, Arizona, U.S.A.

This research analyzes patch development and determines tree spatial patterns along the lower mixed-conifer ecotone on the North Rim of Grand Canyon National Park in Arizona (U.S.A.). Patterns of patch development were interpreted from spatial analyses, based on tree age and size, and past records of disturbance and climate. Five plots in the ecotone between mixed conifer forests and monospecific stands of ponderosa pine (Pinus ponderosa) were studied for patterns of patch development. The methods used include: (1) size-structure analyses, to compare species patch development; (2) dendrochronological dating of tree establishment; (3) tree ring master chronology, to determine periods of suppressed growth, compared to a Palmer Drought Severity Index; and (4) spatial analyses by species composition, size and age, with univariate and bivariate analyses of spatial association and spatial autocorrelation. We found an increased density of shade-tolerant and fire-intolerant species namely clusters of pole-sized white fir, and fewer large ponderosa pine. This revised version was published online in May 2005 with corrections to the Cover Date. This revised version was published online in July 2006 with corrections to the Cover Date.     

Fruit and seed diversity of domesticated carob (Ceratonia siliqua L.) in Morocco

The carob tree (Ceratonia siliqua L.) is an important economic resource for Morocco's rural populations. This species is used in reforestation actions and its cultivation in modern orchards is being undertaken to valorize marginal lands and substitute for drought sensitive species. However, little data is available on its intra-specific variability and its adaptability. Morphological characters of pods and seeds from 13 ecoregions of private-domesticated carob were used to assess phenotypic variation of this species. These stands extend from south-west to north-east and cover a wide range of Morocco's ecoregions. Pods length, width, thickness, seeds number, pulp weight, seeds yield and seeds length, width, thickness and weight were measured for 390 trees (30 trees per ecoregion). Statistically significant differences were found between ecoregions for all characters which were examined, what indicates a high phenotypic diversity. Principal component analysis (PCA) and hierarchical cluster of all ecoregions lead to identify two major and opposite groups (the northern ecoregions; and the central and south-western ecoregions). Ecoregions of the north of Morocco exhibited the largest and the thickest pods with the highest pulp weight while other ecoregions have relatively short pods but largest proportion of seed yield. Similarly, the northern ecoregions are characterized by the heaviest seeds. A correlation matrix between morphological characters, geographic parameters and precipitation exhibits a positive and a negative correlation of pods thickness and pulp weight with the latitude and the altitude, respectively. Seed yield and weight are negatively and positively correlated to pod width, pod thickness and pulp weight, respectively. In addition, seed weight is positively correlated with the latitude. The geographic pattern of the carob tree and its variability are discussed in this paper.     

Multi-scale landscape and seascape patterns associated with marbled murrelet nesting areas on the U.S. west coast

Habitat for wide-ranging species should be addressed at multiple scales to fully understand factors that limit populations. The marbled murrelet (Brachyramphus marmoratus), a threatened seabird, forages on the ocean and nests inland in large trees. We developed statistical relationships between murrelet use (occupancy and abundance) and habitat variables quantified across many spatial scales (statewide to local) and two time periods in California and southern Oregon, USA. We also addressed (1) if old-growth forest fragmentation was negatively associated with murrelet use, and (2) if some nesting areas are more important than others due to their proximity to high quality marine habitat. Most landscapes used for nesting were restricted to low elevation areas with frequent fog. Birds were most abundant in unfragmented old-growth forests located within a matrix of mature second-growth forest. Murrelets were less likely to occupy old-growth habitat if it was isolated (> 5 km) from other nesting murrelets. We found a time lag in response to fragmentation, where at least a few years were required before birds abandoned fragmented forests. Compared to landscapes with little tono murrelet use, landscapes with many murrelets were closer to the ocean's bays, river mouths, sandy shores, submarine canyons, and marine waters with consistently high primary productivity. Within local landscapes (≤ 800ha), inland factors limited bird abundance, but at the broadest landscape scale studied (3200 ha), proximity to marine habitat was most limiting. Management should focus on protecting or creating large, contiguous old-growth forest stands, especially in low-elevation areas near productive marine habitat. This revised version was published online in July 2006 with corrections to the Cover Date.     

Distribution and dynamics of the red squirrel (Sciurus vulgaris L.) in a landscape with fragmented habitat

In a four year study data on the presence of red squirrel were collected in an agricultural landscape by counting dreys in 49 woods ranging from 0.5 to 14 ha, and differing in quality of habitat and isolation.Logit regression analysis showed that the area per woodlot covered with conifers is a good predictor of squirrel presence for each year and during the whole period, but the significance of the regression decreases with time. During the study the number of woods occupied by red squirrel increased, and smaller woods and those without conifers also became inhabited. This trend is in accordance with the positive effect of time in regression analyses on the presence of the species and on the colonization of woods, and it suggests an increase of squirrel numbers in the area. Addition of several isolation variables in the regression analyses showed significant effects in different years, and the effect of isolation was independent of time. In the first two years the area of habitat around a woodlot, the distance to the nearest woodlot larger than 30 ha, and the density of possible movement corridors have significant effects on the presence of red squirrel.In the last two years, with presumably a high number of squirrels, the (short) distance to the nearest woodlot and also the area of habitat around woods have significant effects. It is concluded that the spatial dynamics of the population can be understood as the outcome of individual spatial behaviour, rather than as the result of metapopulation processes.     

Stable Isotope Signatures and Landscape Functioning in Banded Vegetation in Arid-Central Australia

We examined how the measurement of stable carbon and nitrogen isotopes in soils, vegetation and invertebrates can contribute to understanding landscape processes in mulga Acacia aneura ecosystems characterised by alternating wooded groves and intergroves. Our analyses showed that greater leakiness of water from intergroves at the landscape scale tended to promote more conserving physiology at the plant scale. Thus isolated mulga trees in intergroves probably have higher water use efficiency than those in groves. Both trees and grasses in the intergroves have a greater reliance on recycled nitrogen than plants in the groves for which recently fixed N was a substantial source. Grasses in the intergroves had higher N concentrations than those in the groves despite the soil having lower N concentrations. A lack of variation in the isotopic signature of surface soil N suggested that the lower N concentrations in soils of intergroves than groves is due to lower rates of input and not shorter residence times. Stable isotopic signatures of invertebrates showed a diversity of feeding strategies amongst termite species and indicated symbiotic N fixation in two species. There was no relationship between the dependence on N fixation and the habitat preference or diet of termites. Our results suggest that with cautious interpretation, stable isotope signatures could contribute to understanding other ecosystems where patch-interpatch functioning is an important landscape process.     

Estimating regional forest productivity and water yield using an ecosystem model linked to a GIS

We used the PnET-II model of forest carbon and water balances to estimate regional forest productivity and runoff for the northeastern United States. The model was run at 30 arc sec resolution (approximately 1 km) in conjunction with a Geographic Information System that contained monthly climate data and a satellite-derived land cover map. Predicted net primary production (NPP) ranged from 700 to 1450 g m² yr¹ with a regional mean of 1084 g m² yr¹. Validation at a number of locations within the region showed close agreement between predicted and observed values. Disagreement at two sites was proportional to differences between measured foliar N concentrations and values used in the model. Predicted runoff ranged from 24 to 150 cm yr¹with a regional mean of 63 cm yr¹. Predictions agreed well with observed values from U.S. Geologic Survey watersheds across the region although there was a slight bias towards overprediction at high elevations and underprediction at lower elevations.Spatial patterns in NPP followed patterns of precipitation and growing degree days, depending on the degree of predicted water versus energy limitation within each forest type. Randomized sensitivity analyses indicated that NPP within hardwood and pine forests was limited by variables controlling water availability (precipitation and soil water holding capacity) to a greater extent than foliar nitrogen, suggesting greater limitations by water than nitrogen for these forest types. In contrast, spruce-fir NPP was not sensitive to water availability and was highly sensitivity to foliar N, indicating greater limitation by available nitrogen. Although more work is needed to fully understand the relative importance of water versus nitrogen limitation in northeastern forests, these results suggests that spatial patterns of NPP for hardwoods and pines can be largely captured using currently available data sets, while substantial uncertainties exist for spruce-fir.     

Effects of patch shape on the number of organisms

This study examined effects of habitat patch shape on the abundance of organisms. The effects of patch shape were considered in terms of (1) immigration and emigration of organisms. (2) the amount of available resources in a patch and (3) spatial and temporal heterogeneity of the organisms and environment. I hypothesized that (1) the number of organisms would increase as patch shape elongates because organisms are more likely to encounter an elongated patch, (2) the number of organisms in a patch would remain constant for all patch shapes where the number of organisms in a patch was limited by the amount of resources, because patch shape does not change the patch area that is directly associated with the amount of patch resources, and (3) spatial and temporal variation of the abundance of organisms would increase as patch shape elongates because an elongated patch is more likely to interact with the variable surrounding matrix.Common millipedes,Oxidus gracilis, and their habitat, plywood boards of five shapes (width:length ratio; 11, 14, 19, 136, 1144) with an area of 900 cm² were placed in forest and old field and the number of millipedes appearing under the boards was monitored. Significantly higher mean number of millipedes under the boards was observed at a patch with an elongated shape in the forest and the old field. A significant positive correlation was observed between perimeter length of a patch and the number of millipedes in the old field. The temporal and spatial variation of the number of millipedes was high in the old field. The spatial and temporal variation was higher for boards with elongated shape.     

Introducing spatial grazing impacts into the prediction of moorland vegetation dynamics

Grazing by large herbivores is a major determinant of vegetation dynamics in many semi-natural ecosystems, including the replacement of heather moorland by rough grassland in the British uplands. Herbivore foraging is influenced by vegetation patterns and, in turn, their grazing drives vegetation dynamics. Although vegetation impacts are local, spatially heterogeneous local impacts can have different consequences as would the same impacts distributed uniformly. We constructed a simulation model of the spatial effects of grazing by sheep on the vegetation dynamics of heather moorland, a vegetation community of international conservation importance in the UK. The model comprised three sub-models to predict (1) annual average heather utilisation, (2) spatial variation in heather utilisation (higher near the edge of grass patches) and (3) competition between heather and grass. Here we compare the predicted heather utilisation and vegetation dynamics of the spatial model, relative to those of a non-spatial model. The spatial model resulted in a reduced loss of heather cover for a given sheep stocking rate. The model demonstrates how spatial interactions between large herbivores and their forage drive vegetation dynamics, leading to changes in community structure and composition. Indeed, omitting spatial effects in grazing models may lead to inaccurate predictions. We have shown that ecosystem modelling, based around an iterative dialogue between developers and experienced researchers, has the potential to make a substantial contribution towards the conservation and management of vulnerable landscapes. Combining modelling with experimental studies will facilitate progress towards understanding long-term vegetation/herbivore dynamics.     

Introducing spatial grazing impacts into the prediction of moorland vegetation dynamics

Grazing by large herbivores is a major determinant of vegetation dynamics in many semi-natural ecosystems, including the replacement of heather moorland by rough grassland in the British uplands. Herbivore foraging is influenced by vegetation patterns and, in turn, their grazing drives vegetation dynamics. Although vegetation impacts are local, spatially heterogeneous local impacts can have different consequences as would the same impacts distributed uniformly. We constructed a simulation model of the spatial effects of grazing by sheep on the vegetation dynamics of heather moorland, a vegetation community of international conservation importance in the UK. The model comprised three sub-models to predict (1) annual average heather utilisation, (2) spatial variation in heather utilisation (higher near the edge of grass patches) and (3) competition between heather and grass. Here we compare the predicted heather utilisation and vegetation dynamics of the spatial model, relative to those of a non-spatial model. The spatial model resulted in a reduced loss of heather cover for a given sheep stocking rate. The model demonstrates how spatial interactions between large herbivores and their forage drive vegetation dynamics, leading to changes in community structure and composition. Indeed, omitting spatial effects in grazing models may lead to inaccurate predictions. We have shown that ecosystem modelling, based around an iterative dialogue between developers and experienced researchers, has the potential to make a substantial contribution towards the conservation and management of vulnerable landscapes. Combining modelling with experimental studies will facilitate progress towards understanding long-term vegetation/herbivore dynamics.