Overstory-Understory Relationships along Forest Type and Environmental Gradients in the Spring Mountains of Southern Nevada, USA

Isolated forested mountains in deserts have numerous ecological and societal values, but land-management practices (e.g., fire-regime alteration) and climate change can affect forest composition. We analyzed tree overstory-understory relationships on 123 sites in the Spring Mountains within the Mojave Desert near Las Vegas, Nevada, USA to assess three hypotheses. We hypothesized that: the tree species comprising understories are less tolerant of fire than species in overstories, reflecting land-management practices of fire exclusion; mid-elevation forests have the lowest overstory:understory similarity because this zone could have maximum species mixing; and overstory:understory similarity is correlated with environmental gradients (consisting of 14 topographic and soil variables). We found that Pinus monophylla comprised greater relative canopy cover in understories of juniper (32% relative cover) and pinyon-juniper (78%) forests than it did in overstories of these forests (0% and 53%). Similarly, fire-intolerant Abies concolor had 6-fold greater understory than overstory cover in forests with overstories dominated by the fire-tolerant Pinus ponderosa. Overstory:understory S?rensen similarity averaged 43%?77% among six forest types, and there was little support for the supposition that similarity was lowest in mid-elevation forests. Distributions of individual overstory and understory species more closely corresponded with environmental gradients than did overstory:understory similarity. Results suggest that there is high potential for change in at least two of the six dominant forest types of the Spring Mountains. The direction of change (species of moist, higher elevation sites establishing in understories of drier forests) is the opposite of what would be expected for forest adaptation to the warmer, drier, more fire-prone conditions projected for the next century in the southwestern USA.     

Influence of Natal Experience on Nest-Site Selection by Urban-Nesting Cooper's Hawks

Abstract: Exposure to environmental features early in life potentially can influence the kinds of places animals select to live later in life. We examined whether there is evidence that Cooper's hawks (Accipiter cooperii) hatched in an urban environment choose sites with features similar to their natal areas when they nest for the first time. The features we examined were the nest tree species and the level of development surrounding the nest tree. We banded nestling and fledgling Cooper's hawks in Tucson, Arizona, USA, from 1994 to 2004. We then monitored nests in Tucson to identify hawks that had been hatched in the city and eventually secured a breeding site. Percent cover of buildings around first breeding nests was not related to percent cover of buildings around natal nests for either sex. There was some evidence that being hatched in a particular tree species influenced choice of tree species at first breeding sites for males, but the influence was weak. In contrast, tree species in which first-time breeders built their nests, and the sites where the trees were located relative to development, were proportional to what was available in the Tucson metropolitan area. Our data suggest that natal experience played a limited role in nest-site selection by Cooper's hawks in Tucson for the features we examined. If learning occurred, it could have been for the general structure of natal sites. Thus, any small grove of large trees planted in Tucson could be used as a nest site by Cooper's hawks regardless of the level of development surrounding the nest.     

Environmental landscape determinants of maximum forest canopy height of boreal forests

Aims Canopy height is a key driver of forest biodiversity and carbon cycling. Accurate estimates of canopy height are needed for assessing mechanisms relating to ecological patterns and processes of tree height limitations. At global scales forest canopy height patterns are largely controlled by climate, while local variation at fine scales is due to differences in disturbance history and local patterns in environmental conditions. The relative effect of local environmental drivers on canopy height is poorly understood partly due to gaps in data on canopy height and methods for examining limiting factors. Here, we used airborne laser scanning (ALS) data on vegetation structure of boreal forests to examine the effects of environmental factors on potential maximum forest canopy height.     

Detecting Long-term Global Forest Change Using Continuous Fields of Tree-Cover Maps from 8-km Advanced Very High Resolution Radiometer (AVHRR) Data for the Years 1982–99

This paper describes global changes in forest cover from 1982 to 1999 based on the 8-km Pathfinder Advanced Very High Resolution Radiometer (AVHRR) data set. The procedure involves the use of a regression tree in predicting percent tree cover for the years 1982–99. Training data are created from high-resolution imagery and used with phenological metrics derived from the annual AVHRR time series. Using the 18 years of estimated tree cover, and based on a thresholding approach, we identified locations where change in tree cover has occurred. The change sites were then compared to a set of high-resolution deforestation analyses to yield area estimates of deforestation and regrowth. Percent tree cover was found to have decreased globally, from the 1980s to 1990s, in contrast to United Nations Food and Agriculture Organization (FAO) reports of a global increase in forest cover. Latin America and tropical Asia are the two dominant deforestation regions. Paraguay shows the highest rate of forest clearing over the time series, while Indonesia had the greatest increase in deforestation from the 1980s to 1990s. We also suggest that the percent tree-cover maps can be used in standardizing national forest statistics, as an objective means of identifying hot spots of change, and for facilitating ecosystem monitoring.     

Comparison of growth-climate relationship of sabina przewalskii at different timberlines along a precipitation gradient in the northeast Qinghai-Xizang Plateau,China北大核心CSCD

Aims To test the hypothesis that water is the main limiting factor of tree growth at the arid alpine timberline, and to explore the effects of water on growth-climate relationships of Sabina przewalskii along a precipitation gradient in the northeast Qinghai-Xizang Plateau. Methods Three sides were selected to sample the alpine timberline along a precipitation gradient in the northeast Qinghai-Xizang Plateau: Halihatu National Forest Park in Wulan County (HL, annual precipitation 217 mm), Qushigang in Dulan County (QS, 281 mm) and Hebei Forest Farm in Tongde County (HB, 470 mm). The correlation and response analysis at seasonal and extreme climate year scales were used to examine the spatial variations of the growth-climate relationship of S. przewalskii at different timberlines. Important findings Our results do not support the hypothesis that water is the main limiting factor of tree growth at the arid alpine timberline. The effect of precipitation on the radial growth of S. przewalskii were consistent across all three sampling sites, while the effects of temperature were different across sites. At HL site (low precipitation), the winter and summer minimum temperature were the main limiting factor of S. przewalskii radial growth, and this relationship did not significantly change in different extreme climate years. At QS site (middle precipitation), the radial growth of S. przewalskii was mainly limited by the minimum temperature in spring and summer, but its effect was weaker than that at low precipitation site. At HB site (high precipitation), the spring temperature had a significant negative effect on tree growth, and the positive effect of spring precipitation on tree growth was significantly enhanced in comparison with those at low and middle precipitation sites, especially in extreme high temperature and drought years. Summer precipitation did not significantly affect tree growth at high precipitation site. Our results did not support the hypothesis that the radial growth of trees at alpine timberline in arid/humid area is mainly limited by water/temperature. However, precipitation at timberline will affect the relationship between tree growth and temperature at different seasons. With the warming and humidification of the northeastern Qinghai-Xizang Plateau, the climatic limiting factors of tree growth in different timberline areas may be complicated. © 2018 Editorial Office of Chinese Journal of Plant Ecology. All rights reserved.     

Relationships Between Soil Microorganisms, Plant Communities, and Soil Characteristics in Chinese Subtropical Forests

We analyzed the influence of above- and belowground factors on the soil microbial community in a Chinese subtropical forest, one of the most diverse biomes in the northern hemisphere. Soil samples were taken at different depths from four replicate comparative study plots in each of three forest age classes (young 10–40?years, medium 40–80?years, old ≥80?years). Microbial biomass and community structure were then determined using phospholipid fatty acid (PLFA) analysis, and basal respiration and microbial biomass carbon (C_mic) were determined by substrate-induced respiration. These data were then related to plant community and soil variables using non-metric multidimensional scaling analysis and post-hoc permutational correlations. We found that microbial lipid composition and abundance were not related to forest age class. Instead, microbial lipid composition and abundance were related to factors reflecting primary production, i.e., percent litter cover, percent dead wood cover, and percent tree layer cover. Specifically, the relative abundance (mol fraction) of indicators for arbuscular mycorrhizal fungi, Gram-positive and Gram-negative bacteria were positively significantly correlated with percent litter cover. We also found that the biomass of all microbial groups and total PLFA were negatively significantly related to percent deadwood cover. In addition, $ {\text{pH}}_{{{\text{H}}_{ 2} {\text{O}}}} $ was the only soil parameter that was correlated significantly to microbial biomass. Our results indicate that overarching ecological factors such as plant productivity and soil pH are important factors influencing the soil microbial community, both in terms of biomass and of community composition in this subtropical ecosystem.     

Tree diversity,tree height and environmental harshness in eastern and western North America

Does variation in environmental harshness explain local and regional species diversity gradients? We hypothesise that for a given life form like trees, greater harshness leads to a smaller range of traits that are viable and thereby also to lower species diversity. On the basis of a strong dependence of maximum tree height on site productivity and other measures of site quality, we propose maximum tree height as an inverse measure of environmental harshness for trees. Our results show that tree species richness is strongly positively correlated with maximum tree height across multiple spatial scales in forests of both eastern and western North America. Maximum tree height co‐varied with species richness along gradients from benign to harsh environmental conditions, which supports the hypothesis that harshness may be a general mechanism limiting local diversity and explaining diversity gradients within a biogeographic region.     

Subalpine tree growth on serpentine soil: a dendroecological analysis

In this study dendrochronological methods are used to investigate tree growth of several coniferous species growing on serpentine outcrops in the subalpine region. Though stands are thriving on the same parent material, almost barren sites covered with Scots pine trees (Pinus sylvestris/ L.) contrast sharply with adjacent dense stands composed of mixed coniferous species. To find a reasonable explanation for this vegetation pattern and to evaluate the influence of serpentine soil on radial tree growth annual variability in ring-widths was analyzed and compared between both types of stands. Tree-ring analyses show that sparse tree cover on steep slopes might be caused by environmental stress factors especially unfavourable physical soil properties such as instability and dryness. Several abrupt growth reductions followed by slow release in ring-widths indicate repeated mechanical injury to the root system due to soil erosion. On the other hand the aggravating impact of anthropogenic disturbances (tree felling) on this ecologically fragile serpentine area could be determined and dated. There was no evidence that radial tree growth of coniferous species at this site could be limited by toxic heavy metals in the soil.     

Grazing impacts on the diversity and composition of alpine rangelands in Northwest Yunnan

Aims The eastern Himalayan region of Southwest China represents the world's most biodiverse temperate region as well as a cultural hotspot undergoing rapid cultural and ecological change. This area represents the center of origin for many plant groups including horticulturally valuable species of Pedicularis, Rhododendron and Primula. Alpine meadows here also provide summer pastures for Tibetan yak herders and the source for important medicinal plants. Stocking levels for livestock here have quadrupled over the last five decades and shrubs are encroaching into many historical rangelands. Yak herders voice concerns over both shrub encroachment and shrinking grasslands. In this study, we sought to determine: (i) Are alpine rangelands in Deqin County overgrazed and degraded? (ii) What are the local impacts of grazing on plant diversity and community composition? And (iii) which environmental variables covary with these differences in species composition across the grazing gradient?Methods To examine the ecological impacts of grazing in Deqin County, northwest Yunnan and assess its long-term sustainability, we used county records to determine historical population and livestock numbers, confirmed the results of interviews by Haynes (2011) and surveyed plant species richness and cover along a grazing gradient extending away from herder huts. Along these transects, we sampled 1?m 2 quadrats at 5 m intervals, noting species present and percent cover for vascular plants, grasses, sedges, rushes, moss, lichen, exposed rock, bare ground and feces. We also measured the average and maximum plant height within each quadrat. We then computed species richness for each of the 38 transects, calculating alpha and beta diversities. We used one-way ANOVAs to compare mean species richness values and average and maximum plant height across grazing intensities. To chart changes in composition along the 100m gradient, we also plotted the percent cover of graminoid, forb, shrub and bare ground versus distance from the hut. We applied NMS ordination to relate community patterns to environmental variables and grazing intensity using distances to determine species groupings.Important findings Livestock impacts are clearly evident with proportions of grass and bare ground decreasing, shrubs increasing and forbs maintaining even cover with increasing distance from the huts. In comparison with earlier surveys of sites farther from huts, we found reduced plant cover and diversity. Plant species richness almost doubles with increasing distance from herder huts from 9.9 to 19.3 species per 1 m 2 quadrat. An ordination of species and environmental variables demonstrates that grazing strongly affects plant community com position across these plots with strong impacts on palatable plants. If herd sizes remain large and suitable areas for grazing continue to decline, the cumulative impacts of grazing appear likely to degrade the rich diversity of the region and reduce rangeland quality, threatening its ability to sustain current grazing levels.     

Environmental and historical effects on lichen diversity in managed and unmanaged wooded meadows

Question: Which environmental and historical variables affect epiphytic lichen diversity in managed and unmanaged wooded meadows? Location: The island of Gotland located in the Baltic Sea east of the Swedish mainland. Methods: We examined total epiphytic lichen diversity (crustose, foliose and fruticose species) on 1148 trees in eight grazed, eight traditionally managed (mowing, hay gathering and pollarding) and seven unmanaged wooded meadows. In addition to management, data on site location, habitat structure, history and adjacent habitat were analysed. Results: Lichen species richness increased with wooded meadow area and was greater within managed sites than in unmanaged meadows. Historic crown cover (ca. 1930) also influenced present‐day lichen richness. Geographic location, distance to sea, wooded meadow area and average tree circumference were important determinants of lichen species composition. Tree circumference was the strongest overall predictor of the number of species on individual trees. However, tree circumference interacted with management regime, whereby cessation of management appeared to reduce species richness most on large trees. Traditionally managed sites, on average, supported the greatest richness of red‐listed species. Conclusions: Management regime, wooded meadow area and canopy cover were the main drivers of lichen species richness, while geographic location, wooded meadow area, distance to sea and average tree circumference were important determinants of species composition.     

Woodland restoration and forest structure affect nightjar abundance in the Ozark Highlands

Chuck-will's-widow (Antrostomus carolinensis) and eastern whip-poor-will (Antrostomus vociferus) are nightjars in eastern North America that have declined 69% and 67%, respectively, in abundance since 1966, resulting in conservation concerns for these species. We investigated relationships between nightjar abundance and landscape composition, forest structure, and application of tree thinning and prescribed fire because of regional interest in woodland restoration and nightjar conservation. We conducted nocturnal nightjar surveys at 385 points in southern Missouri, USA, in 2014 and 2015 and related counts to pine (Pinus spp.) and hardwood basal area, canopy closure, percent forest cover, and percent of area thinned or burned within 500 m of survey points. We modeled abundance of chuck-will's-widow and eastern whip-poor-will using time-removal models that included a detection process and an abundance process within a hierarchical Bayesian framework. We detected 534 eastern whip-poor-will and 186 chuck-will's-widow during surveys. Our data supported global models that included all 6 vegetation and management variables for both species. Chuck-will's-widow abundance was negatively related to hardwood basal area and peaked at intermediate values of percent area burned and percent forest cover. Eastern whip-poor-will abundance was negatively related to hardwood basal area and canopy cover, positively related to percent forest cover and percent of area burned, and peaked at low to moderate levels of percent of area thinned. Relationships to forest structure and management activities generally supported the conclusion that woodland restoration benefits nightjars and that chuck-will's-widow select landscapes with less forest cover than eastern whip-poor-will.     

Spatial modeling improves understanding patterns of invasive species defoliation by a biocontrol herbivore

Spatial modeling has proven to be useful in understanding the drivers of plant populations in the field of ecology, but has yet to be applied to understanding variation in biocontrol impact. In this study, we employ multi-scale analysis (Moran’s Eigenvector Maps) to better understand the variation in tree canopy exposed to defoliation by a biocontrol beetle (Diorhabda spp.). The control of the exotic tree Tamarix in riparian areas has long been a priority for land managers and ecologists in the American southwest. Diorhabda spp. was introduced as a bio-control agent beginning in 2001 and has since become an inseparable part of Tamarix-dominated river systems in the southwest. Between 2013 and 2016 tamarisk dieback was assessed at 79 sites across Grand County, Utah, arguably the epicenter of Diorhabda impact in the U.S. Canopy cover of Tamarix was between 73 and 81% at these sites, with the percent that was live cover fluctuating by year with a minimum of 42%. Using a traditional general linear model, we found that readily and commonly measured environmental factors could explain only up to 26% of the variation in Tamarix live canopy each year. The number of defoliations was correlated with an increase rather than a decrease in percent live canopy, suggesting compensatory growth. Spatial structure alone explained 22–40% of variation. We found fine scale spatial structure at less than 10 km and broad scale spatial structure from 10 to 30 km. Combining both traditional and novel spatial statistical methods we increased that percentage to 43–63%, depending on year. These results suggest that scientists and land managers must look beyond commonly measured environmental variables to explain non-random biocontrol impact in this system. In particular, this study points to the potential for biotic interactions and variation in flood cycles for further exploration of the identified spatial structure.     

Land‐use legacies in a central Appalachian forest: differential response of trees and herbs to historic agricultural practices

Question: Are contemporary herb and tree patterns explained by historic land use practices? If so, are observed vegetation patterns associated with life‐history characteristics, soil properties, or other environmental variables? Location: Southeastern Ohio, USA. Methods: Using archival records, currently forested sites were identified with distinct land use histories: cultivated, pasture (but not plowed), and reference sites which appear to have never been cleared. Trees were recorded by size and species on twenty 20 m × 20 m plots; percent cover was estimated for each herb species in nested 10 m × 10 m plots. Environmental characteristics were noted, and soil samples analysed for nutrient availability and organic matter. Nonmetric multidimensional scaling ordination was performed separately on both tree and herb datasets to graphically characterize community composition among plots. Life‐history traits were investigated to explain observed compositional differences. Results: Vegetation patterns were explained by current environmental gradients, especially by land‐use history. Cultivated and pasture sites had similar tree composition, distinct from reference sites. Herb composition of pasture and reference sites was similar and distinct from cultivated sites, suggesting the ‘tenacity’ of some forest herbs on formerly cleared sites. Tilling removes rhizomatous species, and disfavors species with unassisted dispersal. These life‐history traits were underrepresented on cultivated sites, although ant‐dispersed species were not. Conclusions: Historic land‐use practices accounted for as much variation in species composition as environmental gradients. Furthermore, trees and herbs responded differently to past land‐use practices. Life‐history traits of individual species interact with the nature of disturbance to influence community composition.     

Recent decline of high altitude coniferous growth due to thermo-hydraulic constrains: evidence from the Miyaluo Forest Reserve,Western Sichuan Plateau of China

Tree growth decline has been reported in many places around the globe under the context of increasingly warming climate, and strengthening drought intensity is detected to be the primary factor for such decline, particularly in northern forest sites, as well as arid and semi-arid areas. Yet, the forest growth decline in high altitude, high mountain sites certainly merits investigation. Here, we reported faxon fir (Abies fargesii var. faxoniana) forest growth decline (slope = -0.64) at the tree line (4150 m above sea level) in Miyaluo Forest Reserve (MFR) at the Western Sichuan Plateau, southwestern China since 2000. We investigated the cause of tree growth decline by applying dendrochronological approaches. We took tree-ring samples from fir trees at the tree line and developed tree-ring width (TRW) chronology. The tree growth – climate relationship analysis showed that maximum temperature (Tmax) was the primary factor limiting the radial growth of fir trees in the investigated area. The moving correlation analysis indicated the strengthening positive influence of Tmax, spring precipitation, and cloud cover during winter and monsoon period on radial growth since 2000s. Our results have shown that both thermal and hydraulic constrains accounted for the radial growth decline of fir trees at the tree line of MFR in the western Sichuan Plateau.     

An Analysis of Overstory Tree Canopy Cover in Sites Occupied by Native and Introduced Cottontails in the Northeastern United States with Recommendations for Habitat Management for New England Cottontail

The New England cottontail (Sylvilagus transitionalis) is a high conservation priority in the Northeastern United States and has been listed as a candidate species under the Endangered Species Act. Loss of early successional habitat is the most common explanation for the decline of the species, which is considered to require habitat with dense low vegetation and limited overstory tree canopy. Federal and state wildlife agencies actively encourage landowners to create this habitat type by clearcutting blocks of forest. However, there are recent indications that the species also occupies sites with moderate overstory tree canopy cover. This is important because many landowners have negative views about clearcutting and are more willing to adopt silvicultural approaches that retain some overstory trees. Furthermore, it is possible that clearcuts with no overstory canopy cover may attract the eastern cottontail (S. floridanus), an introduced species with an expanding range. The objective of our study was to provide guidance for future efforts to create habitat that would be more favorable for New England cottontail than eastern cottontail in areas where the two species are sympatric. We analyzed canopy cover at 336 cottontail locations in five states using maximum entropy modelling and other statistical methods. We found that New England cottontail occupied sites with a mean overstory tree canopy cover of 58% (SE±1.36), and was less likely than eastern cottontail to occupy sites with lower overstory canopy cover and more likely to occupy sites with higher overstory canopy cover. Our findings suggest that silvicultural approaches that retain some overstory canopy cover may be appropriate for creating habitat for New England cottontail. We believe that our results will help inform critical management decisions for the conservation of New England cottontail, and that our methodology can be applied to analyses of habitat use of other critical wildlife species.     

Intercanopy community structure across a heterogeneous landscape in a western juniper‐encroached ecosystem

Questions: How does landscape position influence biotic and abiotic attributes of western juniper (Juniperus occidentalis) encroached ecosystems? How does intercanopy plant community structure respond to changes in soil moisture and temperature based on juniper cover and topographic position? Location: Steens Mountain, southeast Oregon, USA. Objectives: Competition with western juniper modifies plant community composition, alters soil hydrology, and reduces plant productivity. Research is needed to understand these influences across heterogeneous landscapes. This study characterizes the relationship between juniper encroachment and soil water, soil temperature, topographic position, and intercanopy plant community structure. Methods: Using a completely randomized block design, plant density and cover, percent bare ground, percent soil moisture, soil temperature, heat accumulation, and elevation were sampled in 10 m² plots representing low (<1%), moderate (～14%), and high (～27%) juniper cover at four aspects. The relationship and difference between vegetation patterns and environmental variables were analyzed using AOV, NMS, and MRPP (α=0.1). Indicator species analysis tested for shifts in dominant species along ecological gradients. Results: Soil moisture remained higher in low juniper cover sites than moderate and high juniper cover sites. North‐facing sites had highest soil moisture at 5 cm depth with low and moderate juniper cover levels. With increasing soil temperature from May to June, soil moisture declined by 19.7% at 5 cm depth. Achnatherum lemmonii and Pseudoreogneria spicata occurred in closed juniper stands while Achnatherum occidentale and Leymus cinereus were common when encroachment was limited. Application: This approach can be used to predict ecosystem response to western juniper encroachment across heterogeneous landscapes.     

Difference in tree growth responses to climate at the upper treeline: Qilian Juniper in the Anyemaqen Mountains

Three ring-width chronologies were developed from Qilian Juniper (Sabina przewalskii Kom.) at the upper treeline along a west-east gradient in the Anyemaqen Mountains.Most chronological statistics,except for mean sensitivity (MS),decreased from west to east.The first principal component (PC1) Ioadings indicated that stands in a similar climate condition were most important to the variability of radial growth.PC2 Ioadings decreased from west to east,suggesting the difference of tree-growth between eastern and western Anyemaqen Mountains.Correlations between standard chronologies and climatic factors revealed different climatic influences on radial growth along a west-east gradient in the study area.Temperature of warm season (July-August) was important to the radial growth at the upper treeline in the whole study area.Precipitation of current May was an important limiting factor of tree growth only in the western (drier) upper treeline,whereas precipitation of current September limited tree growth in the eastern (wetter) upper treeline.Response function analysis results showed that there were regional differences between tree growth and climatic factors in various sampling sites of the whole study area.Temperature and precipitation were the important factors influencing tree growth in western (drier) upper treeline.However,tree growth was greatly limited by temperature at the upper treeline in the middle area,and was more limited by precipitation than temperature in the eastern (wetter) upper treeline.     

Rates of woody encroachment in African savannas reflect water constraints and fire disturbance

Aim

The aims of this study were to (1) estimate current rates of woody encroachment across African savannas; (2) identify relationships between change in woody cover and potential drivers, including water constraints, fire frequency and livestock density. The found relationships led us to pursue a third goal: (3) use temporal dynamics in woody cover to estimate potential woody cover.

Location

Sub‐Saharan African savannas.

Methods

The study used very high spatial resolution satellite imagery at sites with overlapping older (2002–2006) and newer (2011–2016) imagery to estimate change in woody cover. We sampled 596 sites in 38 separate areas across African savannas. Areas with high anthropogenic impact were avoided in order to more clearly identify the influence of environmental factors. Relationships between woody cover change and potential drivers were identified using linear regression and simultaneous autoregression, where the latter accounts for spatial autocorrelation.

Results

The mean annual change in woody cover across our study areas was 0.25% per year. Although we cannot explain the general trend of encroachment based on our data, we found that change rates were positively correlated with the difference between potential woody cover and actual woody cover (a proxy for water availability; p < .001), and negatively correlated with fire frequency (p < .01). Using the relationship between rates of encroachment and initial cover, we estimated potential woody cover at different rainfall levels.

Main conclusions

The results indicate that woody encroachment is ongoing and widespread across African savannas. The fact that the difference between potential and actual cover was the most significant predictor highlights the central role of water availability and tree–tree competition in controlling change in woody populations, both in water‐limited and mesic savannas. Our approach to derive potential woody cover from the woody cover change trajectories demonstrates that temporal dynamics in woody populations can be used to infer resource limitations.     

Patchiness and co-existence of indigenous and invasive mussels at small spatial scales: the interaction of facilitation and competition

Ecological theory predicts that two species with similar requirements will fail to show long-term co-existence in situations where shared resources are limiting, especially at spatial scales that are small relative to the size of the organisms. Two species of intertidal mussels, the indigenous Perna perna and the invasive Mytilus galloprovincialis, form mixed beds on the south coast of South Africa in a situation that has been stable for several generations of these species, even though these populations are often limited by the availability of space. We examined the spatial structure of these species where they co-exist at small spatial scales in the absence of apparent environmental heterogeneity at two sites, testing: whether conspecific aggregation of mussels can occur (using spatial Monte-Carlo tests); the degree of patchiness (using Korcak B patchiness exponent), and whether there was a relationship between percent cover and patchiness. We found that under certain circumstances there is non-random conspecific aggregation, but that in other circumstances there may be random distribution (i.e. the two species are mixed), so that spatial patterns are context-dependent. The relative cover of the species differed between sites, and within each site, the species with higher cover showed low Korcak B values (indicating low patchiness, i.e. the existence of fewer, larger patches), while the less abundant species showed the reverse, i.e. high patchiness. This relationship did not hold for either species within sites. We conclude that co-existence between these mussels is possible, even at small spatial scales because each species is an ecological engineer and, while they have been shown to compete for space, this is preceded by initial facilitation. We suggest that a patchy pattern of co-existence is possible because of a balance between direct (competitive) and indirect (facilitative) interactions.     

The hydraulic limitation hypothesis revisited

We proposed the hydraulic limitation hypothesis (HLH) as a mechanism to explain universal patterns in tree height, and tree and stand biomass growth: height growth slows down as trees grow taller, maximum height is lower for trees of the same species on resource-poor sites and annual wood production declines after canopy closure for even-aged forests. Our review of 51 studies that measured one or more of the components necessary for testing the hypothesis showed that taller trees differ physiologically from shorter, younger trees. Stomatal conductance to water vapour (g(s)), photosynthesis (A) and leaf-specific hydraulic conductance (K L) are often, but not always, lower in taller trees. Additionally, leaf mass per area is often greater in taller trees, and leaf area:sapwood area ratio changes with tree height. We conclude that hydraulic limitation of gas exchange with increasing tree size is common, but not universal. Where hydraulic limitations to A do occur, no evidence supports the original expectation that hydraulic limitation of carbon assimilation is sufficient to explain observed declines in wood production. Any limit to height or height growth does not appear to be related to the so-called age-related decline in wood production of forests after canopy closure. Future work on this problem should explicitly link leaf or canopy gas exchange with tree and stand growth, and consider a more fundamental assumption: whether tree biomass growth is limited by carbon availability.