The intermediate disturbance hypothesis applies to tropical forests, but disturbance contributes little to tree diversity

The intermediate disturbance hypothesis (IDH) predicts local species diversity to be maximal at an intermediate level of disturbance. Developed to explain species maintenance and diversity patterns in species-rich ecosystems such as tropical forests, tests of IDH in tropical forest remain scarce, small-scale and contentious. We use an unprecedented large-scale dataset (2504 one-hectare plots and 331 567 trees) to examine whether IDH explains tree diversity variation within wet, moist and dry tropical forests, and we analyse the underlying mechanism by determining responses within functional species groups. We find that disturbance explains more variation in diversity of dry than wet tropical forests. Pioneer species numbers increase with disturbance, shade-tolerant species decrease and intermediate species are indifferent. While diversity indeed peaks at intermediate disturbance levels little variation is explained outside dry forests, and disturbance is less important for species richness patterns in wet tropical rain forests than previously thought.     

Impact of pocket gopher disturbance on plant species diversity in a shortgrass prairie community

Summary We examined the impact of pocket gopher disturbances on the dynamics of a shortgrass prairie community. Through their burrowing activity, pocket gophers (Thomomys bottae) cast up mounds of soil which both kill existing vegetation and create sites for colonization by competitively-inferior plant species. Three major patterns emerge from these disturbances: First, we show that 10 of the most common herbaceous perennial dicots benefit from pocket gopher disturbance; that is, a greater proportion of seedlings are found in the open space created by pocket gopher disturbance than would be expected based on the availability of disturbed habitat. Additionally, these seedlings exhibited higher growth rates than adjacent seedlings of the same species growing in undisturbed habitat. Second, we tested two predictions of the Intermediate Disturbance Hypothesis and found that species diversity was greatest for plots characterized by disturbances of intermediate age. However, we did not detect significant differences in diversity between plots characterized by intermediate and high levels of disturbance, indicating that many species are adapted to or at least tolerant of high levels of disturbance. Third, we noted that the abundance of grasses decreased with increasing disturbance, while the abundance of dicots increased with increasing disturbance.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.     

Gap formation and species diversity in Japanese beech forests: a test of the intermediate disturbance hypothesis on a geographic scale

To evaluate whether the intermediate-disturbance hypothesis applies on regional scales, the relationship between the species diversity and gap formation regime of beech forests was examined. The mean gap size and the variation of gap sizes showed no correlation with species diversity. The mean windstorm interval varied widely, but geographical trends, such as latitudinal gradient, were not observed. However, locations that sustained an intermediate frequency of disturbance had the highest species diversity. Although a latitudinal gradient of disturbance was not apparent, the intermediate-disturbance hypothesis was partly supported on a geographic scale. The most predictable model for species diversity was a multiple regression model composed of two factors, the windstorm interval and the cumulative temperature of the growing season. The fact that the temperature was of greater importance than the disturbance interval indicates that the most important factor in predicting forest species diversity is the amount of available energy on a geographic scale.     

The effect of colonization and competition processes on the relation between disturbance and diversity in plant communities

Many theoretical and field studies have emphasized the impact of disturbance in the dynamics and diversity of sessile organism communities. This view is best reflected by the Intermediate Disturbance Hypothesis (IDH), which states that a maximum of diversity is found in ecosystems or communities experiencing intermediate disturbance regimes or at an intermediate stage of development since the last major disturbance event. Although theoretical models based on competitive interactions tend to validate this hypothesis, a recent meta-analysis of field experiments revealed that the mono-modal relationship between disturbance and diversity might not be a general pattern. In this article, we investigate the relationship between disturbance and diversity through the study of patch models, combining two types of competitive interactions: with or without competitive hierarchy, with two mechanisms influencing colonization: negative frequency dependence in colonization rates and immigration. These combinations led to various disturbance-diversity patterns. In the model without competitive hierarchy (founder effect model), a decreasing relationship appeared to be the rule as mentioned in previous studies. In the model with competitive hierarchy, the IDH pattern was obtained for low frequency dependence and low immigration. Nevertheless, high negative frequency dependence in colonization rates led to a decreasing relationship between disturbance and diversity. In contrast, high immigration led to an increasing relationship. The coexistence window (the range of disturbance intensity allowing coexistence) was the widest for intermediate immigration rates. For random species assemblages, patterns with multiple peaks were also possible. These results highlight the fact that the mono-modal IDH pattern should not be considered a rule. Competition and colonization mechanisms have a profound impact on the relationship between disturbance and diversity.     

Dune plant species diversity and function in two barrier island biogeomorphic systems

Barrier island dune systems exhibit strong geographic contrasts in theinteraction between extrinsic disturbance from storm overwash and intrinsicbiogeomorphic recovery processes. To examine how these interactions shape duneplant species diversity, I sampled species cover and topography alongfrequentlystorm-overwashed (South Core Banks, North Carolina) and infrequently overwashed(Sapelo Island, Georgia) barrier islands. The observed compositional anddiversity patterns were in agreement with a complex systems model in whichextrinsic overwash exposure is either reinforced (South Core Banks) or dampened(Sapelo Island) by intrinsic biogeomorphic controls of topography. A largespatial-scale regularity in the distributional pattern of along-shore speciesdiversity was correlated to primary foredune height on South Core. On Sapelo, afine-spatial scale differentiation of species diversity patterns was lessstrongly correlated to topographic metrics. There were no significantdifferences between islands in along-shore alpha diversity (Shannon-Weinerindex). However, Sapelo was more diverse given its smaller area and finer-scalehabitat heterogeneity. I posit that the relevancy of the IntermediateDisturbance Hypothesis is weak when examining diversity patterns along a shoredisturbance gradient. Intrinsic biogeomorphic processes decouple the directcause-and-effect relationship between disturbance and diversity, a basicassumption of IDH. I posit that the Dynamic Equilibrium Model may be a moregenerally applicable conceptual framework. DEM incorporates the interaction ofintrinsic and extrinsic processes that shape habitat heterogeneity, aprerequisite for understanding how complex systems interactions shape diversitypatterns.     

Response of wetland herbaceous communities to gradients of light and substrate following disturbance by thermal pollution

The influence of thermal disturbance and site characteristics on distribution of herbs was studied in portions of a 3020 ha wetland in the southeastern USA. Presence-absence of 52 species in 130 0.25 m² plots was determined from four sites with different disturbance histories and from an undisturbed site. Data from the four disturbed sites were ordinated by detrended correspondence analysis. Differences in species composition among sites (coarse scale) were associated with water depth, light, and substrate type. Within a site (at a fine scale), correlation of environmental variables with ordination scores at a chronically disturbed site was weakly correlated with light (r=0.50). At two sites with episodic disturbance, species composition correlated significantly and positively with substrate and water depth. At a recovering site, vegetation patterns were moderately correlated with water depth (r=?0.52). Species richness was correlated with substrate type along the disturbance gradient. Our results are consistent the intermediate disturbance hypothesis and the subsidy-stress gradient concept.     

The intermediate disturbance hypothesis does not explain fire and diversity pattern in fynbos

The intermediate disturbance hypothesis is a widely accepted generalization regarding patterns of species diversity, but may not hold true where fire is the disturbance. In the Mediterranean-climate shrublands of South Africa, called fynbos, fire is the most importance disturbance and a controlling factor in community dynamics. The intermediate disturbance hypothesis states that diversity will be highest at sites that have had an intermediate frequency of disturbance and will be lower at sites that have experienced very high or very low disturbance frequencies. Measures of diversity are sensitive to scale; therefore, we compared species richness for three fire regimes in South African mountain fynbos to test the intermediate disturbance hypothesis over different spatial scales from 1 m² to 0.1 hectares. Species diversity response to fire frequency was highly scale-dependent, but the relationship between species diversity and disturbance frequency was opposite that predicted by the intermediate disturbance hypothesis. At the largest spatial scales, species diversity was highest at the least frequently burned sites (40 years between fires) and lowest at the sites of moderate (15 to 26 years between fires) and high fire frequency (alternating four and six year fire cycle). Community heterogeneity, measured both as the slope of the species-area curve for a site and as the mean dissimilarity in species composition among subplots within a site, correlated with species diversity at the largest spatial scales. Community heterogeneity was highest at the least frequently burned sites and lowest at the sites that experienced an intermediate fire frequency.     

Species diversity,susceptibility to disturbance and tree population dynamics in tropical rain forest

Abstract. In 1964 a census of all trees > 9.7 cm diameter at breast height (DBH) was conducted on 22 plots totalling 13.2 ha in lowland tropical evergreen rain forest on Kolombangara, Solomon Islands. Over the following 30 yr (1964–1994), populations of all individuals > 4.85 cm DBH of the 12 most common tree species and amounts of disturbance have been monitored on a declining number of these plots (in 1994, nine plots totalling 5.4 ha were still being recorded). Between November 1967 and April 1970, Kolombangara was struck by four cyclones, although only two of these caused substantial amounts of damage to the canopy structure. Multivariate analysis has identified six forest types on Kolombangara (Greig-Smith et al. 1967). The species richness and diversity of trees in the 1964 census, turnover rates of the populations monitored over 1964–1975, and the amount of disturbance sustained during a cyclone in 1970, were all positively correlated across five of the forest types. The sixth forest type was a consistent outlier in these analyses and is believed to have been seriously disturbed by humans about a century ago. The floristics, turnover and disturbance data support Connell's intermediate disturbance hypothesis. The most species-rich forest types contained a higher proportion of fast-growing individuals and species that are early-successional and which have low density timber. Properties of these species rendered them more susceptible to damage when struck by the 1970 cyclone. They showed higher turnover rates because disturbance-dependent species are also characterised by higher mortality and recruitment rates. Thus, periodic cyclones appear to favour the maintenance of differences in species diversity and composition between forest types.     

Interspecific variation in tree seedling establishment in canopy gaps in relation to tree density

Abstract. We tested whether interspecific variation in tree seedling establishment in canopy gaps was significantly related to interspecific variation in tree density, for seven deciduous forest tree species (Quercus alba, Hamamelis virginiana, Acer rubrum, Sassafras albidum, Quercus rubra, Prunus serotina, Ostrya virginiana). For each species, seedling establishment was calculated as the difference in seedling density before experimental gap creation versus three years after gap creation. In each of the six experimentally-created gap types (33 % or 66 % removal of tree basal area from 0.01-ha, 0.05-ha or 0.20-ha patches), differences in seedling establishment among species were significantly related to differences in their density in the tree canopy. A regression model with log_e tree density as the independent variable accounted for between 93 % and 98 % of interspecific variation in seedling establishment. Our results provide empirical support for models of tree dynamics in gaps that assume seedling establishment depends on canopy tree density.     

Tree species richness complementarity, disturbance and fragmentation in a Mexican tropical montane cloud forest

Tropical montane cloud forest exhibits great heterogeneity in speciescomposition and structure over short geographic distances. In central Veracruz,Mexico, the conservation priority of seven cloud forest fragments was assessedby considering differences in woody plant species richness and complementarityof species among sites, forest structure, tree mortality, and timber andfirewood extraction as indicators of anthropogenic disturbance. Densities oftrees 5 cm dbh (360–1700 trees/ha) weredifferent among the sites, but basal area (35.3–89.3m²/ha) did not differ among fragments. The number of dead trees rangedfrom 10–30 to 170–200 trees/ha. The fragments'species composition was different but complementary. The Morisita–Hornindex indicated low similarity between fragments. A non-parametric estimator ofspecies richness indicated that more sampling effort would be necessary tocomplete the inventory (15 additional trees and two understory shrub species).Unfortunately, most of the fragments are threatened with deforestation. The numberof cut trees was similar among sites (0–15 cut trees/0.1 ha).Sites with immediate need for conservation were close to settlements, with highnumbers of cut trees and no legal protection. The selected sites represent thevariety of situations that exist in the region. Given the high complementarityobserved between fragments, a regional conservation approach will be required topreserve the last repositories of part of the tremendous biodiversity of theonce continuous forest in this region.     

Effects of tree species diversity on a community of weaver spiders in a tropical forest plantation

The effects of producer diversity on predators have received little attention in arboreal plant communities, particularly in the tropics. This is particularly true in the case of tree diversity effects on web‐building spiders, one of the most important groups of invertebrate predators in terrestrial plant communities. We evaluated the effects of tree species diversity on the community of weaver spiders associated with big‐leaf mahogany (Swietenia macrophylla) in 19, 21 × 21‐m plots (64 plants/plot) of a tropical forest plantation which were either mahogany monocultures (12 plots) or polycultures (seven plots) that included mahogany and three other tree species. We conducted two surveys of weaver spiders on mahogany trees to evaluate the effects of tree diversity on spider abundance, species richness, diversity, and species composition associated with mahogany. Our results indicated that tree species mixtures exhibited significantly greater spider abundance, species richness, and diversity, as well as differences in spider species composition relative to monocultures. These results could be due to species polycultures providing a broader range of microhabitat conditions favoring spider species with different habitat requirements, a greater availability of web‐building sites, or due to increased diversity or abundance of prey. Accordingly, these results emphasize the importance of mixed forest plantations for boosting predator abundance and diversity and potentially enhancing herbivore pest suppression. Future work is necessary to determine the specific mechanisms underlying these patterns as well as the top‐down effects of increased spider abundance and species richness on herbivore abundance and damage.     

Vegetation analysis and tree population structure of tropical wet evergreen forests in and around Namdapha National Park,northeast India

Species composition, diversity and tree population structure were studied in three stands of the tropical wet evergreen forest in and around Namdapha National Park, Arunachal Pradesh, India. Three study stands exposed to different intensities of disturbances were identified, viz., undisturbed (2.4 ha) in the core zone of the park, moderately disturbed (2.1 ha) in the periphery of the park and highly disturbed (2.7 ha) outside the park area. In total 200 plant species belonging to 73 families were recorded in three stands. Tree density and basal area showed a declining trend with the increase in disturbance intensity. The densities of tree saplings and seedlings were lower in the disturbed stands than in the undisturbed stand. Species like Altingia excelsa, Olea dioica, Terminalia chebula, Mesua ferrea and Shorea assamica in the undisturbed stand and Albizia procera alone in the moderately disturbed stand contributed more than 50% of the total tree density in respective stands. The undisturbed stand contained young tree population. In the highly disturbed stand, the tree density was scarce, but had uncut trees of higher girth class (>210 cm GBH). Low shrub density was recorded in both disturbed stands due to frequent human disturbances; the broken canopy and direct sunlight enhanced the abundance of herbs in these stands. With a species rarity (species having <2 individuals) of ca. 50%, the tropical wet evergreenforests of the Namdapha National Park and its adjacent areas warrant more protection from human intervention and also eco-development to meet the livelihood requirements of the local inhabitants in the peripheral areas of the Namdapha National Park in order to reduce the anthropogenic pressure on the natural resources of the park.     

Long-term observations of rain forest succession,tree diversity and responses to disturbance.

The relationship between succession and tropical forest diversity hasbeen much debated. A fundamental disagreement hinges on whether high localspecies richness is a transient successional property, albeit one that can bemaintained by disturbance, or is rather a property of stable late successionalcommunities. This paper addresses this controversy employing a series oflong-term permanent sample plot data spanning seven decades. W.J. Eggelingstudied the vegetation of Budongo Forest, Uganda during the 1930s and 1940s. Hedescribed a series of ten plots (1.4 and 1.86 ) as asuccessionalprogression of forest types in which tree species numbers show a unimodalrise-and-fall over time – a pattern best known from Connell'sillustration of his intermediate disturbance hypothesis. Tree communities infive of the original plots have been intermittently re-assessed over thesubsequent decades. One data-series provides observations spanning 54-yearsfromone intact undisturbed old-growth forest plot. The remaining fourplots were assessed before and after controlled disturbances (tree poisoning)executed in the late 1950s and early 1960s, and the resulting data-series spanc. 20 years of pre-disturbance and c.35 years of post-disturbance changes. Theunimodal pattern of species-richness in the original comparative plot-series isparalleled by a similar rise-and-fall in stem-densities, but rarefactionconfirms that the unimodal pattern in richness also holds for fixedstem-counts.The proportion of species occurring in both large and small stem-size-classesincreases across the series. As richness declines in later succession, lowabundance species occur predominantly in larger stem-sizes. All time-seriesshowa rise in species richness ranging from 12 to 177% (over 50–60 years). Each of thedisturbed plots ultimately reaches greater richness than was recorded anywherein Eggeling's original series. Contrary to expectation a small rise wasalso recorded in the undisturbed late successional plot (c.42 species 10 diameter ha^–1, rising to c.47). The lowestspecies density observed in the study is a 1940s record of c. 10 species 10 diameter ha^–1 in monodominantCynometra [Caesalpinoidae] forest and thehighestrecord is c. 61 recorded in 1992, in theyoungest vegetation type monitored. These observations indicate both thevolatile nature of tree-richness patterns and the limitations of simple modelsas aids to interpretation when confronted with real patterns of long-termchange.     

The response of herbaceous-layer vegetation to anthropogenic disturbance in intermittent stream bottomland forests of southern Indiana,USA

Between 1993 and 1995 we sampled herbaceous layer vegetation on 84 plots in Platanus/Asarum Wet-Mesic Bottomland forests to determine how these forests have responded to human disturbance. Four different disturbance types were sampled (abandoned agricultural are as, clearcuts, group-selection openings, and single-tree selection openings), and uncut 80–100 year-old reference stands were sampled for comparison. Detrended Correspondence Analysis (DCA), distance analyses (chord distance and normalized Euclidean distance) and similarity analysis (Bray-Curtis similarity coefficient) suggest that agricultural use has shifted herbaceous-layer vegetation composition away from that typical of the reference forests, but that clearcutting, group-selection harvest, and single-tree selection harvest have not greatly shifted herbaceous composition. This shift in vegetation on abandoned agricultural land resulted from a loss of indicator species, such as Cardamine concatenata (Michx.) Sw., Stellaria pubera Michx., and Laportea canadensis (L.) Weddell and an influx of disturbance, exotic, and nonforest species (e.g., Lycopodium complanatum L., Lonicera japonica Thunb. and Senecio aureus L.). However, only two species found in reference stands, Erigenia bulbosa (Michx.) Nutt. and Sphenopholis obtusata (Michx.) Scribn., were missing from clearcuts, group-selection openings, and single-tree selection openings. The species richness values of abandoned agriculture, clearcut, and group-selection plots were generally greater than those of single-tree selection and reference plots. Abandoned agricultural areas had much greater total species richness because of the influx of dry-site, exotic, disturbance, and non-forest species.     

Herbaceous species responses to long-term effects of prescribed fire, grazing and selective tree cutting in the savanna-woodlands of West Africa

Grazing, fire and selective tree cutting are major disturbances that shape species diversity in savanna ecosystems, yet their effects are highly variable. We carried out a factorial experiment with two levels to examine the effects of grazing, fire and selective tree cutting on herbaceous species richness, abundance and diversity on two sites in the Sudanian savanna-woodlands of Burkina Faso for 10 years (1994–2003). The results showed significant inter-annual variation in species richness, abundance and diversity at both sites (p<0.001), while main or combined effects of fire, grazing and selective cutting were very limited and varied between life forms and sites. Grazing tended to favour the diversity of perennial grasses; fire tended to influence the richness of annual grasses and abundance and diversity of perennial grasses while selective tree cutting had no effect on any of the vegetation attributes assessed. The combined effect of grazing, fire and selective cutting tended to increase the diversity of forbs. In many cases, the responses of herbaceous species to treatments were clearer on the site with deeper soils than the one with shallow soils. Depending on the site and treatments, the inter-annual variation in vegetation attributes was partly related to amount and/or frequency of rainfall and partly to inter-annual variation in grazing or fire intensity. It can be concluded that both disturbances and climatic condition influence the structure and diversity of herbaceous flora in the Sudanian savanna-woodland ecosystem. The responses were site-specific, which accentuates the importance of landscape-scale approaches to understand the impacts of disturbances on composition, structure and diversity of savanna ecosystems.