Scale dependence of vegetation-environment correlations: A case study of a North Carolina piedmont woodland

Abstract. Vegetation and its correlation with environment has been traditionally studied at a single scale of observation. If different ecological processes are dominant at different spatial and temporal scales, the results obtained from such observations will be specific to the single scale of observation employed and will lack generality. Consequently, it is important to assess whether the processes that determine community structure and function are similar at different scales, or whether, how rapidly, and under what circumstances the dominant processes change with scale of observation. Indeed, early work by Greig-Smith and associates (Greig-Smith 1952; Austin & Greig-Smith 1968; see Greig-Smith 1979; Kershaw & Looney 1985; Austin & Nicholls 1988) suggested that plant-plant interactions are typically important at small scales, but that the physical environment dominates at large scales. Using a gridded and mapped 6.6 ha portion of the Duke Forest on the North Carolina piedmont for a case study, we examined the importance of scale in vegetation studies by testing four hypotheses. First, we hypothesized that the correlation between vegetation composition and environment should increase with increasing grain (quadrat) size. Our results support this hypothesis. Second, we hypothesized that the environmental factors most highly correlated with species composition should be similar at all grain sizes within the 6.6-ha study area, and should be among the environmental factors strongly correlated with species composition over the much larger extent of the ca. 3500 ha Duke Forest. Our data are not consistent with either portion of this hypothesis. Third, we hypothesized that at the smaller grain sizes employed in this study (< 256 m²), the composition of the tree canopy should contribute significantly to the vegetation pattern in the under-story. Our results do not support this hypothesis. Finally, we predicted that with increased extent of sampling, the correlation between environment and vegetation should increase. Our data suggest the opposite may be true. This study confirms that results of vegetation analyses can depend greatly on the grain and extent of the samples employed. Whenever possible, sampling should include a variety of grain sizes and a carefully selected sample extent so as to ensure that the results obtained are robust. Application of the methods used here to a variety of vegetation types could lead to a better understanding of whether different ecological processes typically dominate at different spatial scales.     

High-elevation rock outcrop vegetation of the Southern Appalachian Mountains

Abstract. Species composition patterns and vegetation-environment relationships were quantified for high-elevation rock outcrops of the Southern Appalachian Mountains, an infrequent and insular habitat in a forested landscape. Outcrops occur over a wide geographic range encompassing extensive variation in both geology and climate. Geographic-scale factors interact with site-scale factors to produce variation in vegetation among outcrops. Similarly, site-scale factors interact with micro-scale factors to produce variation in vegetation within outcrops. To provide a quantitatively-based classification of outcrop vegetation we used a TWINSPAN analysis of 154 100-m² plots. We recognized nine communities that primarily correspond to different combinations of elevation, bedrock type, geography, and moisture. Within outcrops of a single bedrock type, vegetation composition of 100-m² plots was consistently correlated with elevation and solar radiation, but relationships to soil nutrients varied with bedrock type. Both site-scale (100 m²) factors (e.g. elevation, slope, aspect, and bedrock type) and plot-scale (1-m²) microsite factors (e.g. soil depth, vegetation height, soil nutrients) were strongly correlated with species composition at the 1-m² level. Environment can be used to predict composition more effectively for 100-m² plots on a single bedrock type than either across bedrock types or at a 1-m² scale. Composition-environment relationships resemble those described for outcrop systems from other regions with pronounced topographic relief more than they do those described for the nearby but flatter and lower-elevation outcrops of the Southeastern Piedmont. There is strong spatial autocorrelation in this community, perhaps owing to dispersal limitation. Consequently, a comprehensive conservation strategy must include reservation of both a range of geologic types and a range of geographic locations.     

Vegetation change following removal of keystone herbivores from desert grasslands in New Mexico

Abstract. Responses of plant communities to mammalian herbivores vary widely, due to variation in plant species composition, herbivore densities, forage preferences, soils, and climate. In this study, we evaluated vegetation changes on 30 sites within and adjacent to the Sevilleta National Wildlife Refuge (SNWR) in central New Mexico, USA, over a 20‐yr period following removal of the major herbivores (livestock and prairie dogs) in 1972–1975. The study sites were established in 1976, and were resampled in 1986 and 1996 using line transect methods. At the landscape scale, repeated measures ANOVA of percentage cover measurements showed no significant overall net changes in total perennial plant basal cover, either with or without herbivores present; however, there was an overall increase in annual forbs and plant litter from 1976 to 1996. At the site scale, significant changes in species composition and dominance were observed both through time and across the SNWR boundary. Site histories varied widely, with sites dominated by Bouteloua eriopoda being the most dynamic and sites dominated by Scleropogon brevifolius being the most persistent. Species‐specific changes also were observed across multiple sites: B. eriopoda cover increased while Gutierrezia sarothrae greatly decreased. The non‐uniform, multi‐directional changes of the sites' vegetation acted to prevent detection of overall changes in perennial vegetation at the landscape level. Some sites displayed significant changes after removal of herbivores, while others appeared to respond primarily to climate dynamics. Certain species that were not preferred by livestock or prairie dogs, showed overall declines during drought periods, while other preferred species exhibited widespread increases during wetter periods regardless of herbivore presence. Therefore, the vegetation dynamics cannot be attributed solely to removal of herbivores, and in some cases can be explained by short‐ and long‐term fluctuations in climate. These results emphasize the variety of responses of sites with differences in vegetation to mammalian herbivores under otherwise similar climatic conditions, and illustrate the value of site‐ and landscape‐scale approaches to understanding the impacts of plant‐herbivore interactions.     

Does diversity beget diversity? A case study of crops and weeds

Abstract. The ecological literature is ambiguous as to whether the initial diversity of a plant community facilitates or deters the diversity of colonizing species. We experimentally planted annual crop species in monoculture and polyculture, and examined the resulting weed communities. The species composition of weeds was similar among treatments, but the species richness of weeds was significantly higher in the polycultures than in the monocultures. This supports the ‘diversity begets diversity’ hypothesis. Environmental microheterogeneity, diversity promoters, and ecological equivalency do not seem able to explain the observed patterns.     

The effects of treefall gaps on understory vegetation in New York State

Abstract. Treefalls are a common form of disturbance in northeastern United States forests. The resultant gaps contribute to a high degree of environmental heterogeneity in the understory of these forests. Plant density, plant cover, and species richness in understory plant communities were monitored for three years during the growing season, May - September. Differences between treefall gap and closed canopy vegetation were less pronounced early in the growing season for plant density and leaf cover. Species richness was significantly greater within treefall gaps during the entire growing season. Eight species were found in greater abundance within treefall gaps (i.e., gap-phase species), while one species was found more commonly under closed canopy. Ordination results suggest that time since gap creation and treefall gap size marginally affect the species composition of vegetation found within treefall gaps.     

Phytogeographical and community similarities of alpine tundras of Changbaishan Summit,China, and Indian Peaks,USA

Abstract. We compared the diversity, phytogeography, and plant communities in two mid-latitude alpine tundras with comparable aerial and elevational extents: Changbaishan Summit in eastern Asia and Indian Peaks in western North America. Despite wide separation, the two areas shared 72 species. In all, 43% of the species on Changbaishan Summit are also distributed in the alpine zones of western North America, while 22% of the species on Indian Peaks are also distributed in the alpine zones of eastern Asia. Almost all the shared species also occur in the Beringian region. Phytogeographical profiles of species and genera showed that 69% of species and over 90% of genera in both alpine tundras belong to the three phytogeographical categories: cosmopolitan, circumpolar, and Asian-North American. We attributed the current floristic relationship between these widely separated areas to the periodic past land connection between the two continents during the Tertiary and Pleistocene. Indian Peaks has a closer floristic relationship with the Arctic tundra than does Changbaishan Summit. Indian Peaks also has 45% higher species richness and lower vegetation cover than Changbaishan Summit. Plant communities from the two areas were completely separated in the two-way indicator species analysis and non-metric multidimensional scaling on floristic data at both species and generic levels, whereas ordination of communities by soil data produced a greater overlap. The plant communities on Changbaishan Summit in general have lower alpha diversity, higher beta diversity (lower between-community floristic similarity), and more rare species than does Indian Peaks. Mosaic diversity does not differ in the two alpine tundras, although the analysis suggests that Changbaishan Summit communities are more widely spaced on gradients than the Indian Peaks communities.     

How fast is the carousel? Direct indices of species mobility with examples from an Oklahoma grassland

Abstract. Current interest in small‐scale species dynamics has led to a proliferation of mobility indices. We advocate the use of direct measures of mobility such as immigration rate, extinction rate, residence time, and carousel time. We also demonstrate that the null expectation of cumulative frequency under different null models can be calculated explicitly. Species can depart from the commonly‐used ‘random reassignment’ model simply because of longevity, and not mobility per se. We therefore prefer a random immigration null model, which assumes that immigration locations are randomized. We examined mobility patterns of selected plant species, studied in 256 quadrats of each of four grains (ranging from 1/64 m² to 1 m²) in an Oklahoma grassland. Residence times and carousel times can be centuries or even millennia for some species. We explore the numerical and biological reasons for relationships between mobility statistics. Mobility statistics are fairly consistent among grains and years, although the residence times of species exhibit some subtle scale dependence. Species depart from a random immigration model very slightly – but the departure is consistent: species tend to re‐occupy previously vacated space more often than expected due to chance. We believe that the use of direct indices will facilitate the study of how species characteristics influence mobility.     

Predicting vegetation types at treeline using topography and biophysical disturbance variables

Abstract. The relationships between four vegetation types and variables representing topography and biophysical disturbance gradients were modeled for a study area in east-central Glacier National Park, Montana. Four treeline transition vegetation types including closed-canopy forest, open-canopy forest, meadow, and unvegetated surfaces (e.g. rock, snow, and ice) were identified and mapped through classification of satellite data and subsequent field verification. Topographic characteristics were represented using a digital elevation model and three variables derived from topoclimatic potential models (solar radiation potential, snow accumulation potential, and soil saturation potential). A combination of generalized additive and generalized linear modeling (GAM and GLM, respectively) techniques was used to construct logistic regression models representing the distributions of the four vegetation types. The variables explained significant amounts of variation in the vegetation types, but high levels of variation remained unexplained. A comparison of ‘expected’ and ‘observed’ vegetation patterns suggested that some unexplained variation may have occurred at the basin scale. A suite of tools and techniques is presented that facilitates predicting landscape-scale vegetation patterns and testing hypotheses about the spatial controls on those patterns.     

Fine root biomass within experimental canopy gaps: evidence for a below-ground gap

To study fine root biomass in response to canopy gap formation, we determined fine root biomass in the upper 15 cm of soil within four experimental canopy gaps, and within the surrounding intact forest. Sampling was conducted throughout the growing season (May through August) following gap creation. We observed a mid-season decrease in root biomass within gaps, and within the intact forest surrounding the gaps during June and July. Later in the season, less fine root biomass was found within canopy gaps than within the intact forest, and this trend may be attributed to root death accompanying the tree(s) that were felled to form the gaps. These results provide strong evidence for the formation of a below-ground gap in association with the creation of a canopy gap.     

Vegetation patterns in heterogeneous landscapes: The importance of history and environment

Abstract. Throughout the eastern United States, plant species distributions and community patterns have developed in response to heterogeneous environmental conditions and a wide range of historical factors, including complex histories of natural and anthropogenic disturbance. Despite increased recognition of the importance of disturbance in determining forest composition and structure, few studies have assessed the relative influence of current environment and historical factors on modern vegetation, in part because detailed knowledge of prior disturbance is often lacking. In the present study, we investigate modern and historical factors that control vegetation patterns at Harvard Forest in central Massachusetts, USA. Similar to the forested uplands throughout the northeastern United States, the site is physiographically heterogeneous and has a long and complex history of natural and anthropogenic disturbance. However, data on forest composition and disturbance history collected over the past > 90 years allow us to evaluate the importance of historical factors rigorously, which is rarely possible on other sites. Soil analyses and historical sources document four categories of historical land use on areas that are all forested today: cultivated fields, improved pastures/mowings, unimproved pastures, and continuously forested woodlots. Ordination and logistic regressions indicate that although species have responded individualistically to a wide range of environmental and disturbance factors, many species are influenced by three factors: soil drainage, land use history, and C:N ratios. Few species vary in accordance with ionic gradients, damage from the 1938 hurricane, or a 1957 fire. Contrary to our expectation that the effects of disturbance will diminish over time, historical land use predicts 1992 vegetation composition better than 1937 composition, perhaps because historical woodlots have become increasingly differentiated from post-agricultural stands through the 20th century. Interpretations of modern vegetation must consider the importance of historical factors in addition to current environmental conditions. However, because disturbances such as land use practices and wind damage are complex, it is often difficult to detect disturbance effects using multivariate approaches, even when the broad history of disturbance is known.