Variation in Susceptibility to Hurricane Damage as a Function of Storm Intensity in Puerto Rican Tree Species

One of the most significant challenges in developing a predictive understanding of the long-term effects of hurricanes on tropical forests is the development of quantitative models of the relationships between variation in storm intensity and the resulting severity of tree damage and mortality. There have been many comparative studies of interspecific variation in resistance of trees to wind damage based on aggregate responses to individual storms. We use a new approach, based on ordinal logistic regression, to fit quantitative models of the susceptibility of a tree species to different levels of damage across an explicit range of hurricane intensity. Our approach simultaneously estimates both the local intensity of the storm within a plot and the susceptibility to storm damage of different tree species within plots. Using the spatial variation of storm intensity embedded in two hurricanes (Hugo in 1989 and Georges in 1998) that struck the 16 ha Luquillo Forest Dynamics Plot in eastern Puerto Rico, we show that variation in susceptibility to storm damage is an important aspect of life history differentiation. Pioneers such as Cecropia schreberiana are highly susceptible to stem damage, while the late successional species Dacryodes excelsa suffered very little stem damage but significant crown damage. There was a surprisingly weak relationship between tree diameter and the susceptibility to damage for most of the 12 species examined. This may be due to the effects of repeated storms and trade winds on the architecture of trees and forest stands in this Puerto Rican subtropical wet forest.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp .     

Discontinuous DBH–height relationship of <Emphasis Type="Italic">Cryptomeria japonica</Emphasis> on Yakushima Island: effect of frequent typhoons on the maximum height

The mountain zone of Yakushima Island is covered with a mixed conifer-broadleaved forest dominated by old-growth Cryptomeria japonica (L.f.) D. Don trees. Even though Yakushima Island has been frequently struck by typhoons with wind velocities exceeding 55 m s⁻¹, the Cr. japonica trees in the mountain zone have survived for thousands of years without fatal damage. To evaluate the effect of storms on tree growth, the relationships between the stem diameter at breast height (DBH) and the heights of Cr. japonica and coexistent tree species were investigated. Two models based on an expanded allometric equation and a discontinuous piecewise allometric equation, respectively, to represent DBH–height relationships were evaluated. In all plots, the DBH–height relationship of Cr. japonica was discontinuous between small DBH and large DBH individuals. The tops of the large DBH individuals of Cr. japonica were lost to strong winds. However in each instance, they occupied the highest position in the canopy, even if they had lost their tops. In contrast, the DBH–height relationships of subcanopy broadleaved species were continuous in many plots and the equilibrium heights of the dominant broadleaved species were similar and almost in the same order regardless of the canopy heights of Cr. japonica. These results revealed a constant vertical structure in the Cr. japonica forest on Yakushima Island. Our results demonstrate a vertical niche segregation in the forest under high wind pressures and such vertical structure enables effective use of forest space and increases the basal area density.     

Changes in two Minnesota forests during 14 years following catastrophic windthrow

Abstract. We measured tree damage and mortality following a catastrophic windthrow in permanent plots in an oak forest and a pine forest in central Minnesota. We monitored changes in forest structure and composition over the next 14 years. Prior to the storm, the oak forest was dominated by Quercus ellipsoidalis, and the pine forest by Pinus strobus. The immediate impacts of the storm were to differentially damage and kill large, early‐successional hardwoods and pines. Subsequent recovery was characterized by the growth of late‐successional hardwoods. In both forests the disturbance acted to accelerate succession. Ordination of tree species composition confirmed the trend of accelerated succession, and suggested a convergence of composition between the two forests.     

Vegetation dynamics in central european forest ecosystems (near-natural as well as managed) after storm events

All over the world forests and woodlands are damaged or reset to initial stages by fire, insect outbreaks or storms. In Central Europe storm events are the most important natural disturbances affecting stand structures of both natural and managed forests and yet only a few studies exist on long-term forest development following the destruction of the tree layer by a storm. This paper presents a permanent plot study established in 1988 in the Bavarian Forest National Park (SE Germany) on areas, where the tree layer had been destructed by a storm on August 1, 1983. The records concerning (1) floristic composition (spermatophytes, pteridophytes, bryophytes, lichens) and cover degree, (2) location and shape of each tree higher than 1 meter (height, diameter at breast height) including position of fallen trees and (3) number of seedlings and saplings were taken in 1988, 1993 and 1998. Two windfall areas, situated next to each other in the same broad and flat valley bottom on wet soils under local cold climate conditions (potential as well as recent vegetation:Calamagrostio villosae-Piceetum bazzanietosum) were analyzed, one of them with completely free development after the storm event (“untouched”), the other with dead wood cleared off after the event, but thereafter with free development (“cleared”). The vegetation analysis separated two major trends in vegetation dynamics: (1) On the cleared plots with intensive soil-surface disturbance (removal of the damaged wood) the species composition changed towards pioneer herb vegetation (Rubus sp.), and pioneer forest species (here: birch,Betula pendula and/orB. pubescens) established. Subsequently, vegetation dynamics leading towards clusters of forest ground-layer species composition took place. (2) In untouched stands, where soil-surface disturbances were restricted to pit-and-mound-system created by uprooted trees, the patchiness of forest vegetation increased and a regeneration of mainly terminal tree species (here: Norway spruce,Picea abies) started. Stand development for the next 100 years was simulated using the model FORSKA-M. The model is individual-based and includes competition for light, soil water, and nutrients. The simulations suggest that floristic structures of cleared and untouched plots, respectively, will remain different for several decades, but within one century, the floristic structure becomes rather similar. Major processes in forest ecosystems which can be used to improve forest management and nature conservation practices have been identified based on the results of the case study.     

Linking responses of native and invasive plants to hurricane disturbances: implications for coastal plant community structure

Hurricane disturbances produce significant changes in forest microclimates, creating opportunities for seedling regeneration of native and invasive plant species alike. However, there is limited information on how changes in microclimates and pre-existing forest conditions affect native and invasive plants responses to hurricane disturbances. In this manipulative study, we examined the responses of three common shrub/small stature tree species, two of which are native to the coastal region of the southeastern USA (Baccharis halimifolia and Morella cerifera) and one that is invasive (Triadica sebifera), to two key components of hurricane disturbance (canopy damage and saline storm surge). In a greenhouse, we grew seedlings of these species under a range of shade levels that mimicked pre-and post-hurricane canopy conditions for wet pine forest and mixed hardwood forest, two forest communities common in coastal areas of the southeastern USA. Seedlings were subjected to saline storm surges equivalent to full strength sea water for 3 days. Seedling responses (mortality and growth) to the treatments were monitored for 16 months. All species benefitted from higher canopy openness. Storm surge effects were short-lived and seedlings readily recovered under high light conditions. The storm surge had stronger negative effects on survival and growth of all species when coupled with high shade, suggesting storm surge has greater negative impacts on seedlings where hurricane winds cause minimal or no canopy damage. The invasive T. sebifera was by far more shade tolerant than the natives. Survival of T. sebifera seedlings under highly shaded conditions may provide it a competitive edge over native species during community reassembly following tropical storms. Differential responses of native and invasive species to hurricane disturbances will have profound consequences on community structure across coastal forest stands, and may be regulated by legacies of prior disturbances, community structure, extent of canopy damage, and species’ tolerance to specific microclimates.

     

Pervasive growth reduction in Norway spruce forests following wind disturbance

Background

In recent decades the frequency and severity of natural disturbances by e.g., strong winds and insect outbreaks has increased considerably in many forest ecosystems around the world. Future climate change is expected to further intensify disturbance regimes, which makes addressing disturbances in ecosystem management a top priority. As a prerequisite a broader understanding of disturbance impacts and ecosystem responses is needed. With regard to the effects of strong winds – the most detrimental disturbance agent in Europe – monitoring and management has focused on structural damage, i.e., tree mortality from uprooting and stem breakage. Effects on the functioning of trees surviving the storm (e.g., their productivity and allocation) have been rarely accounted for to date.

Methodology/Principal Findings

Here we show that growth reduction was significant and pervasive in a 6.79·million hectare forest landscape in southern Sweden following the storm Gudrun (January 2005). Wind-related growth reduction in Norway spruce (Picea abies (L.) Karst.) forests surviving the storm exceeded 10% in the worst hit regions, and was closely related to maximum gust wind speed (R² = 0.849) and structural wind damage (R² = 0.782). At the landscape scale, wind-related growth reduction amounted to 3.0 million m³ in the three years following Gudrun. It thus exceeds secondary damage from bark beetles after Gudrun as well as the long-term average storm damage from uprooting and stem breakage in Sweden.

Conclusions/Significance

We conclude that the impact of strong winds on forest ecosystems is not limited to the immediately visible area of structural damage, and call for a broader consideration of disturbance effects on ecosystem structure and functioning in the context of forest management and climate change mitigation.     

Hurricane disturbance in a temperate deciduous forest: patch dynamics, tree mortality, and coarse woody detritus

Patch dynamics, tree injury and mortality, and coarse woody detritus were quantified to examine the ecological impacts of Hurricane Fran on an oak-hickory-pine forest near Chapel Hill, NC. Data from long-term vegetation plots (1990–1997) and aerial photographs (1998) indicated that this 1996 storm caused patchy disturbance of intermediate severity (10–50% tree mortality; Woods, J Ecol 92:464–476, 2004). The area in large disturbance patches (>0.1 ha) increased from <1% to approximately 4% of the forested landscape. Of the forty-two 0.1-ha plots that were studied, 23 were damaged by the storm and lost 1–66% of their original live basal area. Although the remaining 19 plots gained basal area (1–15% increase), across all 42 stands basal area decreased by 17% because of storm impacts. Overall mortality of trees >10 cm dbh was 18%. The basal area of standing dead trees after the storm was 0.9 m²/ha, which was not substantially different from the original value of 0.7 m²/ha. In contrast, the volume and mass of fallen dead trees after the storm (129 m³/ha; 55 Mg/ha) were 6.1 and 7.9 times greater than the original levels (21 m³/ha; 7 Mg/ha), respectively. Uprooting was the most frequent type of damage, and it increased with tree size. However, two other forms of injury, severe canopy breakage and toppling by other trees, decreased with increasing tree size. Two dominant oak species of intermediate shade-tolerance suffered the largest losses in basal area (30–41% lost). Before the storm they comprised almost half of the total basal area in a forest of 13% shade-tolerant, 69% intermediate, and 18% shade-intolerant trees. Recovery is expected to differ with respect to vegetation (e.g., species composition and diversity) and ecosystem properties (e.g., biomass, detritus mass, and carbon balance). Vegetation may not revert to its former composition; however, reversion of biomass, detritus mass, and carbon balance to pre-storm conditions is projected to occur within a few decades. For example, the net change in ecosystem carbon balance may initially be negative from losses to decomposition, but it is expected to be positive within a decade after the storm. Repeated intermediate-disturbance events of this nature would likely have cumulative effects, particularly on vegetation properties.     

Modelling the wind damage probability in forests in Southwestern Germany for the 1999 winter storm ‘Lothar’

The wind damage probability (P _DAM) in the forests in the federal state of Baden-Wuerttemberg (Southwestern Germany) was calculated using weights of evidence (WofE) methodology and a logistic regression model (LRM) after the winter storm ‘Lothar’ in December 1999. A geographic information system (GIS) was used for the area-wide spatial prediction and mapping of P _DAM. The combination of the six evidential themes forest type, soil type, geology, soil moisture, soil acidification, and the ‘Lothar’ maximum gust field predicted wind damage best and was used to map P _DAM in a 50 × 50 m resolution grid. GIS software was utilised to produce probability maps, which allowed the identification of areas of low, moderate, and high P _DAM across the study area. The highest P _DAM values were calculated for coniferous forest growing on acidic, fresh to moist soils on bunter sandstone formations—provided that ‘Lothar’ maximum gust speed exceeded 35 m s⁻¹ in the areas in question. One of the most significant benefits associated with the results of this study is that, for the first time, there is a GIS-based area-wide quantification of P _DAM in the forests in Southwestern Germany. In combination with the experience and expert knowledge of local foresters, the probability maps produced can be used as an important tool for decision support with respect to future silvicultural activities aimed at reducing wind damage. One limitation of the P _DAM-predictions is that they are based on only one major storm event. At the moment it is not possible to relate storm event intensity to the amount of wind damage in forests due to the lack of comprehensive long-term tree and stand damage data across the study area.     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

Ice‐storm disturbance and long‐term forest dynamics in the Adirondack Mountains

Ice storms cause periodic disturbance to temperate forests of eastern North America. They are the primary agents of disturbance in some eastern forests. In this paper, a forest gap model is employed to explore consequences of ice storms for the long‐term dynamics of Tsuga canadensis‐northem hardwoods forests. The gap model LINKAGES was modified to simulate periodic ice storm disturbance in the Adirondack Mountains of New York. To adapt the gap model for this purpose, field data on ice storm disturbance are used to develop a polytomous logistic regression model of tree damage. The logistic regression model was then incorporated into the modified forest gap model, LINK ADIR, to determine the type of damage sustained by each simulated tree. The logistic regression model predicts high probabilities of bent boles or severe bole damage (leaning, snapping, or uprooting) in small‐diameter trees, and increasing probability of canopy damage as tree size increases. Canopy damage is most likely on gentle slopes; the probability of severe bole damage increases with increasing slope angle. In the LINKADIR simulations, tree damage type determines the probability of mortality; trees with severe bole damage are assigned the highest mortality rate. LINKADIR predicts Tsuga canadensis dominance in mesophytic old‐growth forests not disturbed by ice storms. When ice storms are simulated, the model predicts Acer saccharum‐dominated forests with higher species richness. These results suggest that ice storms may function as intermediate disturbances that enhance species richness in forested Adirondack landscapes.     

Hyperspectral remote detection of niche partitioning among canopy trees driven by blowdown gap disturbances in the Central Amazon

Advanced recruitment and neutral processes play important roles in determining tree species composition in tropical forest canopy gaps, with few gaps experiencing clear secondary successional processes. However, most studies are limited to the relatively limited spatial scales provided by forest inventory plots, and investigations over the entire range of gap size are needed to better understand how ecological processes vary with tree mortality events. This study employed a landscape approach to test the hypothesis that tree species composition and forest structural attributes differ between large blowdown gaps and relatively undisturbed primary forest. Spectral mixture analysis on hyperspectral satellite imagery was employed to direct field sampling to widely distributed sites, and blowdown plots were compared with undisturbed primary forest plots. Tree species composition and forest structural attributes differed markedly between gap and non-gap sites, providing evidence of niche partitioning in response to disturbance across the region. Large gaps were dominated by classic Neotropical pioneer genera such as Cecropia and Vismia, and average tree size was significantly smaller. Mean wood density of trees recovering in large gaps (0.55 g cm⁻³) was significantly lower than in primary forest plots (0.71 g cm⁻³), a difference similar to that found when comparing less dynamic (i.e., tree recruitment, growth, and mortality) Central Amazon forests with more dynamic Western Amazon forests. Based on results, we hypothesize that the importance of neutral processes weaken, and niche processes strengthen, in determining community assembly along a gradient in gap size and tree mortality intensity. Over evolutionary time scales, pervasive dispersal among colonizers could result in the loss of tree diversity in the pioneer guild through competitive exclusion. Results also underscore the importance of considering disturbance processes across the landscape when addressing forest carbon balance.     

Long-term changes in a Wisconsin Fagus-Acer forest in relation to glaze storm disturbance

Changes in the composition of a Fagus-Acer (Beech-Sugar maple) forest in southeastern Wisconsin over a 16-yr period from 1971 to 1987 are analyzed in relation to a severe glaze (ice) storm disturbance occurring within the census period. Landscape topography created ‘windward’ and ‘leeward’ forest aspects with respect to storm severity, which resulted in greater canopy opening on the windward aspect. In the tree stratum, most species remained stable in density and most of the common species increased in basal area into larger size classes. However, Fagus grandifolia, Ulmus rubra, and the small tree Ostrya virginiana suffered net losses that suggest synergistic effects between glaze storm disturbance and other factors upon tree mortality. In the sapling stratum, canopy opening strongly promoted release of shade-to levant Acer sac-charum. On the windward forest aspect, sapling densities of less shade-tolerant species also increased, in contrast to the absence of such increases on the leeward forest aspect. In the shrub (regeneration) stratum, species responses were heterogeneous. Regeneration of most species increased over the 16-yr period, and some less shade-tolerant species showed increased regeneration differentially on the windward forest aspect. Overall, disturbance appears to have accelerated forest succession toward increased dominance by A. saccharum and persistence of both Fagus and Tilia americana through their capacities for root sprouting. However, forest succession was retarded somewhat on the windward aspect through increased recruitment of less-shade tolerant species. These results parallel those of other studies of glaze storm disturbance, and they illustrate how spatially heterogeneous disturbance intensity may contribute to maintenance of forest diversity.     

大兴安岭不同演替阶段天然林物种多度格局模拟

为实现大兴安岭地区落叶松天然林的全周期可持续经营,以新林林业局翠岗林场3块100m×100m不同演替阶段(白桦林、落叶松-白桦混交林和落叶松林)的天然林固定调查样地数据为基础,采用统计模型(对数正态模型)、生态位模型(断棍模型、生态位优先占领模型、Zipf模型、Zipf-Mandelbrot模型)和中性模型(群落零和多项式模型、Volkov模型)对不同演替阶段天然林的乔木层、幼苗层和由乔木层划分的不同林层的物种多度格局进行拟合,并用χ² 检验和Kolmogorov-Smirnov检验选择最佳模型。结果表明:(1) 在大兴安岭地区不同演替阶段的落叶松天然林群落中,Zipf-Mandelbrot模型的拟合效果均最好。(2) 对数正态模型对于稳定的演替阶段白桦林和落叶松林的群落相较于过渡阶段群落落叶松-白桦混交林的物种多度分布的拟合效果更好。(3) 在大兴安岭地区落叶松天然林中,生境过滤主导群落构建,随着群落不断演替生境过滤和中性过程减弱。(4) 天然林群落乔木层的群落构建过程与乔木层内不同生长阶段林层的构建过程不同,生境过滤过程在落叶松林乔木层中逐渐减弱,而在落叶松林不同生长阶段的林层和更新层中增强。     

外来树种日本落叶松和日本花柏对鄂西北森林群落的影响

【目的】研究大面积栽植外来树种日本落叶松和日本花柏对鄂西北森林群落物种多样性的影响,为评价外来树种的入侵风险提供依据。【方法】采用样地调查法对比研究外来树种日本落叶松和日本花柏人工林与当地森林群落特征差异。【结果】日本落叶松人工林样地林下植物种类丰富,样地平均物种数与当地天然林样地物种数相比无显著差异;林下灌木层和草本层多样性指数较高,草本层多样性指数与当地天然林相比无显著差异;林下未发现日本落叶松的更新苗。日本花柏人工林样地平均物种数显著少于当地其他人工林样地;乔木层、灌木层和草本层的4个多样性指数值均小于当地其他人工林样地;在3个样地中出现日本花柏更新苗,更新苗数量达479株·hm~(-2)。【结论】引种栽培日本落叶松不会导致当地森林群落的物种多样性降低,不会引起生物入侵风险;但引种栽培日本花柏会明显降低当地森林群落的物种多样性,对当地森林生态系统的稳定构成一定威胁。     

Spatial patterns and interspecific associations of dominant tree species in two old-growth karst forests,SW China

Spatial patterns and interspecific associations of plant species in forests are important for revealing how species interact with each other and with the environment, and hence have important implications for optimal forest management and restoration in degraded forest ecosystems. In this paper, the O-ring statistics were used to characterize the spatial patterns and interspecific associations of eight dominant tree species in two 1-ha old-growth karst forest plots in Maolan National Natural Reserve, southwestern China. We found that most of the eight dominant tree species in two forests were continuously regenerating populations. Six species (Platycarya longipes, Acer wangchii, Clausena dunniana, Castanopsis carlesii var. spinulosa, Distylium myricoides, and Rhododendron latoucheae) exhibited significant aggregations at the majority of scales while others (Celtis biondii and Cyclobalanopsis myrsinaefolia) showed a random distribution pattern at most scales. Negative association was a dominant pattern for most species pairs in the two plots, while positive associations were found at most scales for only two species pairs (Platycarya–Clausena and Castanopsis–Rhododendron). Results also indicated that the two main factors of habitat complexity and heterogeneity—the elevation and rock-bareness rate—play important roles in determining spatial distribution patterns and interspecific associations of tree species in karst forests of Maolan. Thus, the observed spatial patterns among the eight tree species are influenced by habitat heterogeneity in the context of karst topographical variations. The partitioning of habitat niches contributes to the promoting species coexistence in species-rich karst forests. The differences of species features in spatial patterns and associations should be paid more attention when planning forest management and developing restoration strategies.     

Position on slope,disturbance, and tree species coexistence in a Seasonal Semideciduous Forest in SE Brazil

We investigated the influence of position on a slope (plot relative elevation) and vegetation disturbance (the tallest tree height per plot) on community composition and diversity in a SE Brazilian Seasonal Semideciduous Forest (46°55′ W, 22°50′ S). Trees with dbh ≥5 cm were sampled in one hundred 10 × 10 m plots randomly placed in a 6.5-ha stand. Through partial Mantel test, floristic dissimilarities among plots (Jaccard index computed with species abundance in each plot) were correlated with environmental distances among plots (Euclidian distance index computed with relative elevation and the tallest tree height values in each plot). Relative elevation and the tallest tree per plot height were individually correlated with floristic gradients expressed by PCA axes scores using Pearson’s correlation coefficient. Through resampling, we compared diversity (richness, Berger-Parker D and Shannon H′) among plots in the drier (up) and moister (low) ends of the slope. Floristic dissimilarities were significantly correlated with environmental distances even after geographic distances among plots have been partialled out (r _m = 0.1274, p < 0.001). The first two PCA axes accounted for 22% of the total variance. After Bonferroni and Dutilleul’s corrections, axis 1 showed a marginally significant correlation with plot relative elevation (r = − 0.4097, p = 0.0309), and axis 2 was significantly correlated with the tallest tree height per plot (r = 0.2953, p = 0.0106). Position on the slope and vegetation disturbance were reliable predictors of community composition, thus suggesting the operation of niche assembly organizing processes. Richness and diversity (H′) decreased and dominance (D) increased with elevation on the slope. Dominance increase from D ₍₃₀₀₎ = 0.11 (confidence interval = 0.091–0.131) to D ₍₃₀₀₎ = 0.19 (CI = 0.165–0.210) surpassed the expected dominance increase based on the reduction of richness alone: D ₍₃₀₀₎ = 0.13 (CI = 0.110–0.140), thus highlighting the niche partitioning assembly of the community, especially among abundant species. Given the great amount of floristic variability remaining unexplained, stochastic processes, such as those related to dispersal limitation, may also have influence on the community composition. Therefore, both niche assembly and chance events can operate even on a fine local scale.     

The decline of tree diversity on newly isolated tropical islands: A test of a null hypothesis and some implications

Summary Six islands, each less than a hectare in area, were isolated in about 1913 from the mainland of central Panamá by the rising waters of Gatun Lake. By 1980, the diversity of trees on all but one of these islands was far lower than on mainland plots of comparable size. A restricted subset of tree species has spread on these islands, notablyProtium panamense, Scheelea zonensis, Oenocarpus panamanus andSwartzia simplex. We constructed a null model to predict how chance would change tree diversity and the similarity of tree species compositions of different islands, assuming that each mature tree has equal chances of dying and/or reproducing, regardless of its species. This model cannot account for the diminished diversity of the changes in vegetation on these islands: some factors must be favoring a particular set of tree species.Two factors, exposure to wind and absence of mammals, seem needed to bring about the vegetation changes observed on these small islands. Their vegetation shows many signs of wind damage and of adaptation to resist wind, reflecting its exposure to dry season winds and storm winds sweeping across the lake from the west. Their most common tree species appear to have spread because mammals rarely visit these small and isolated islands. Seed of these common species are normally much eaten by mammals and do not need burial by mammals to escape insect attack.A thorough grasp of plant—animal interactions is needed to understand the events that have taken place on these islands. Identifying those keystone animals essential for maintaining plant diversity is a necessary element of reserve design and forest management in the tropics.The US government has the right to retain a non-exclusive, royalty-free license in and to any copyright covering this paper.     

How selective are elephants as agents of forest tree damage in Bwindi Impenetrable National Park,Uganda?

Elephants are locally concentrated in Bwindi Impenetrable National Park. Vegetation damage attributable to elephants appears to be increasing and may result in the modification of the forest. We examined the implied selectivity of stem damage due to elephants. We followed 26.84 km of recent elephant trails and used 122 plots to document tree damage in relation to species, stem sizes and locations. Of 897 trees (DBH ≥2 cm), 542 (60.4%) were intact, 22 (2.5%) debarked, 274 (30.5%) toppled and 172 (19.2%) had broken branches. Small trees were more likely to be pushed over or have their branches broken, whereas large trees were more commonly debarked. The species most frequently selected for damage included mid‐successional species such as Newtonia buchananii, Myrianthus holstii and Chrysophyllum albidum. These species may be vulnerable to increasing elephant numbers. Our analyses using general linear models indicate that elephants are selective concerning where, how and what tree stems they damage. We found a higher incidence of elephant damage per‐tree stem in open areas than in more closed areas, suggesting feedback in which elephants maintain open habitats that may be conducive for other species such as mountain gorillas. More work is needed to better determine how changing elephant numbers may influence Bwindi's conservation values.     

Sapling herbivory,invertebrate herbivores and predators across a natural tree diversity gradient in Germany’s largest connected deciduous forest

Tree species-rich forests are hypothesised to be less susceptible to insect herbivores, but so far herbivory–diversity relationships have rarely been tested for tree saplings, and no such study has been published for deciduous forests in Central Europe. We expected that diverse tree communities reduce the probability of detection of host plants and increase abundance of predators, thereby reducing herbivory. We examined levels of herbivory suffered by beech (Fagus sylvatica L.) and maple saplings (Acer pseudoplatanus L. and Acer platanoides L.) across a tree species diversity gradient within Germany’s largest remaining deciduous forest area, and investigated whether simple beech or mixed stands were less prone to damage caused by herbivorous insects. Leaf area loss and the frequency of galls and mines were recorded for 1,040 saplings (>13,000 leaves) in June and August 2006. In addition, relative abundance of predators was assessed to test for potential top-down control. Leaf area loss was generally higher in the two species of maple compared to beech saplings, while only beech showed a decline in damage caused by leaf-chewing herbivores across the tree diversity gradient. No significant patterns were found for galls and mines. Relative abundance of predators on beech showed a seasonal response and increased on species-rich plots in June, suggesting higher biological control. We conclude that, in temperate deciduous forests, herbivory–tree diversity relationships are significant, but are tree species-dependent with bottom-up and top-down control as possible mechanisms. In contrast to maple, beech profits from growing in a neighbourhood of higher tree richness, which implies that species identity effects may be of greater importance than tree diversity effects per se. Hence, herbivory on beech appeared to be mediated bottom-up by resource concentration in the sampled forest stands, as well as regulated top-down through biocontrol by natural enemies. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorised users.     

Non-native grass invasion suppresses forest succession

Multiple factors can affect the process of forest succession including seed dispersal patterns, seedling survival, and environmental heterogeneity. A relatively understudied factor affecting the process of succession is invasions by non-native plants. Invasions can increase competition, alter abiotic conditions, and provide refuge for consumers. Functional traits of trees such as seed size and life history stage may mediate the effects of invasions on succession. We tested the effects of the forest invader Microstegium vimineum on planted and naturally regenerating trees in a multi-year field experiment. We established plots containing nine species of small- and large-seeded tree species planted as seeds or saplings, and experimentally added Microstegium to half of all plots. Over 3 years, Microstegium invasion had an overall negative effect on small-seeded species driven primarily by the effect on sweetgum, the most abundant small-seeded species, but did not affect large-seeded species such as hickory and oak species, which have more stored seed resources. Natural regeneration was over 400% greater in control than invaded plots for box elder, red maple, and spicebush, and box elder seedlings were 58% smaller in invaded plots. In contrast to the effects on tree seedlings, invasion did not affect tree sapling survival or growth. Microstegium may be directly reducing tree regeneration through competition. Invaded plots had greater overall herbaceous biomass in 2006 and 2008 and reduced light availability late in the growing season. Indirect effects may also be important. Invaded plots had 120% more thatch biomass, a physical barrier to seedling establishment, and significantly greater vole damage to tree saplings during 2006 and 2007. Our results show that two tree functional traits, seed size and life history stage, determined the effects of Microstegium on tree regeneration. Suppression of tree regeneration by Microstegium invasions may slow the rate of forest succession and alter tree species composition.