Species phylogenetic relatedness, priority effects, and ecosystem functioning

Species immigration history can structure ecological communities through priority effects, which are often mediated by competition. As competition tends to be stronger between species with more similar niches, we hypothesize that species phylogenetic relatedness, under niche conservatism, may be a reasonable surrogate of niche similarity between species, and thus influence the strength of priority effects. We tested this hypothesis using a laboratory microcosm experiment in which we established bacterial species pools with different levels of phylogenetic relatedness and manipulated the immigration history of species from each pool into microcosms. Our results showed that strong priority effects, and hence multiple community states, only emerged for the species pool with the greatest phylogenetic relatedness. Community assembly also resulted in a significant positive relationship between bacterial phylogenetic diversity and ecosystem functions. Interestingly, these results emerged despite a lack of phylogenetic conservatism for most of the bacterial functional traits considered. Our results highlight the utility of phylogenetic information for understanding the structure and functioning of ecological communities, even when phylogenetically conserved functional traits are not identified or measured.     

Interacting effects of habitat destruction and changing disturbance rates on biodiversity: Who is going to survive?

Changes in disturbance rates due to climate change may increase or decrease diversity, whereas permanent loss of habitat is generally believed to decrease diversity. It is, however, very likely that the effects of disturbances and habitat destruction interact. Understanding such combined effects is essential to predict the response of communities to global changes and in particular which functional types of species are most endangered. Using an individual-based spatially explicit community model, we investigate (1) whether diversity-disturbance curves alter when spatially uncorrelated or autocorrelated habitat destruction is added, and (2) which functional types of species are able to survive under these altered conditions. Model communities consisted of four functional types of species trading off between colonisation ability and competition strength. We found that habitat destruction may alter both height and shape of diversity-disturbance curves: maximum diversity at intermediate disturbance rates may shift to other disturbance rates or even split into two peaks giving rise to bimodal diversity-disturbance relationships with different sub-communities persisting at low and high disturbance rates. Diversity responded differentially depending on how the colonisation-competition trade-off was represented. Our results suggest that, for trade-offs in seed production rate, generally the best coloniser will better withstand the interacting effects of habitat destruction and changing disturbance rates; however, for trade-offs in mean dispersal distances, functional types characterized by intermediate abilities will perform best. We conclude that predictions of the impacts of changing disturbance rates on biodiversity depend on community structure and cannot be made without knowledge of concurrent permanent habitat destruction.     

Scale-Dependent Effects of Habitat Disturbance on Species Richness in Tropical Forests

Abstract: Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.     

Community assembly in experimental grasslands: suitable environment or timely arrival?

It is hard to defend the view that biotic communities represent a simple and predictable response to the abiotic environment. Biota and the abiotic environment interact, and the environment of an individual certainly includes its neighbors and visitors in the community. The complexity of community assembly calls forth a quest for general principles, yet current results and theories on assembly rules differ widely. Using a grassland microcosm as a model system, we manipulated fertility, disturbance by defoliation, soil/microclimate, and arrival order of species belonging to two groups differing in functional attributes. We analyzed the outcome of community assembly dynamics in terms of species richness, invasibility, and species composition. The analyses revealed strong environmental control over species richness and invasibility. Species composition was mainly determined by the arrival order of species, indicating that historical contingency may change the outcome of community assembly. The probability for multiple equilibria appeared to increase with productivity and environmental stability. The importance of arrival order offers an explanation of the difficulties in predicting local occurrences of species in the field. In our experiment, variation in fertility and disturbance was controlling colonization with predictable effects on emergent community properties such as species richness. The key mechanism is suggested to be asymmetric competition, and our results show that this mechanism is relatively insensitive to the species through which it works. While our analyses indicate a positive and significant correlation between richness and invasibility, the significance disappears after accounting for the effect of the environment. The importance of arrival order (historical contingency) and environmental control supports the assumption of the unified neutral theory that different species within a trophic level can be considered functionally equivalent when it comes to community assembly. However, our results indicate that variation in asymmetric competition is the key factor determining the richness of the resulting communities, and this is far from neutral.     

Impacts of chronic trawling disturbance on meiofaunal communities

Bottom trawling causes chronic and widespread disturbance to the seabed in shelf seas. Meiofauna may be impacted directly or indirectly by this disturbance, since the passage of trawls causes immediate mortality or displacement, changes sediment structure and geochemistry and affects the abundance of predators or competitors. Since meiofauna make a significantly greater contribution to benthic production than the larger macrofauna, there are compelling reasons to assess their response to chronic trawling disturbance. In this study, we determined the effects of trawling disturbance, season, sediment type and depth on the structure and diversity of nematode communities. Our analyses were based on comparisons between nematode communities in three beam-trawl fishing areas in the central North Sea. These areas were trawled with mean frequencies of I (low disturbance), 4 (medium) and 6 (high) times year⁻¹ respectively. Our analyses showed that trawling had a significant impact on the composition of nematode assemblages. In two sampling seasons, the number of species, diversity and species richness of the community were significantly lower in the area subject to high levels of trawling disturbance than in the areas subject to low or medium levels of disturbance. However, levels of disturbance at the ‘low’ and ‘medium’ sites may have been insufficient to cause marked longterm changes in community structure. Many of the observed changes were consistent with responses to other forms of physical disturbance. The extent to which the observed changes in community structure reflect changes in the production of the nematode community remains unknown, although overall abundance was not significantly affected by trawling disturbance. Published online. 9 August 2002     

Disturbance, Diversity, and Invasion: Implications for Conservation

Disturbance is an important component of many ecosystems, and variations in disturbance regime can affect ecosystem and community structure and functioning. The "intermediate disturbance hypothesis" suggests that species diversity should be highest at moderate levels of disturbance. However, disturbance is also known to increase the invasibility of communities. Disturbance therefore poses an important problem for conservation management, Here, we review the effects of disturbances such as fire grazing, soil disturbance and nutrient addition on plant species diversity and invasion with particular emphasis on grassland vegetation. Individual components of the disturbance regime can have marked effects on species diversity, but it is often modifications of the existing regime that have the largest influence. Similarly, disturbance can enhance invasion of natural communities, but frequently it is the interaction between different disturbances that has the largest effect. The natural disturbance regime is now unlikely to persist within conservation areas since fragmentation and human intervention have usually modified physical and biotic conditions. Active management decisions must now be made on what disturbance regime is required and this requires decisions on what species are to be encouraged or discouraged.     

Organization of Plethodon salamander communities: guild-based community assembly

A long-standing goal in evolutionary ecology is to determine whether the organization of communities is reflective of underlying deterministic processes. In this study, I examined patterns of species co-occurrence among eastern Plethodon salamanders and determined whether they were consistent with predictions from a guild model of competition-based community assembly. Using a database of 45 species and 4540 geographic sites, I found that patterns of co-occurrence were significantly nonrandom at both a regional and continental scale, and species of different size guilds were distributed more evenly in sites than was expected by chance. Sites with the highest species richness had consistent patterns of community composition, and with few exceptions, the same five species were present at all sites. Taken together, these results imply that larger Plethodon communities are assembled from simpler communities in a manner consistent with what is predicted through competitive mechanisms and suggest that stable species combinations are possible to achieve at various levels of species richness. These results also provide strong evidence consistent with the hypothesis that competitive-based community assembly is a general phenomenon in Plethodon and that interspecific competition is prevalent among the eastern species of this group.     

Perturbations alter community convergence, divergence, and formation of multiple community states

Environmental perturbations (e.g., disturbance, fertilization) commonly shift communities to a new mean state, but much less is known about their effects on the variability (dispersion) of communities around the mean, particularly when perturbations are combined. Community dispersion may increase or decrease (representing a divergence or convergence among communities) if changing environmental conditions alter species interactions or magnify small initial differences that develop during community assembly. We used data from an experimental study of disturbance and fertilization in a low-productivity grassland to test how these two perturbations affect patterns of species composition and abundance. We found that a one-time biomass reduction decreased community dispersion, which persisted over four growing seasons. Conversely, continuous fertilization increased community dispersion and, when combined with disturbance, led to the formation of three distinct community states. These results illustrate that perturbations can have differing effects on community dispersion. Attention to the variance in community responses to perturbations lends insight into how ecological interactions determine community structure, which may be missed when focusing only on mean responses. Furthermore, multiple perturbations may have complex effects on community dispersion, yielding convergence or divergence patterns that are difficult to predict based on analysis of single factors.     

Diversity-invasibility across an experimental disturbance gradient in Appalachian Forests

Research examining the relationship between community diversity and invasions by nonnative species has raised new questions about the theory and management of biological invasions. Ecological theory predicts, and small-scale experiments confirm, lower levels of nonnative species invasion into species-rich compared to species-poor communities, but observational studies across a wider range of scales often report positive relationships between native and nonnative species richness. This paradox has been attributed to the scale dependency of diversity-invasibility relationships and to differences between experimental and observational studies. Disturbance is widely recognized as an important factor determining invasibility of communities, but few studies have investigated the relative and interactive roles of diversity and disturbance on nonnative species invasion. Here, we report how the relationship between native and nonnative plant species richness responded to an experimentally applied disturbance gradient (from no disturbance up to clearcut) in oak-dominated forests. We consider whether results are consistent with various explanations of diversity-invasibility relationships including biotic resistance, resource availability, and the potential effects of scale (1 m2 to 2 ha). We found no correlation between native and nonnative species richness before disturbance except at the largest spatial scale, but a positive relationship after disturbance across scales and levels of disturbance. Post-disturbance richness of both native and nonnative species was positively correlated with disturbance intensity and with variability of residual basal area of trees. These results suggest that more nonnative plants may invade species-rich communities compared to species-poor communities following disturbance.     

Dominant species identity, not community evenness, regulates invasion in experimental grassland plant communities

While there has been extensive interest in understanding the relationship between diversity and invasibility of communities, most studies have only focused on one component of diversity: species richness. Although the number of species can affect community invasibility, other aspects of diversity, including species identity and community evenness, may be equally important. While several field studies have examined how invasibility varies with diversity by manipulating species identity or evenness, the results are often confounded by resource heterogeneity, site history, or disturbance. We designed a mesocosm experiment to examine explicitly the role of dominant species identity and evenness on the invasibility of grassland plant communities. We found that the identity of the dominant plant species, but not community evenness, significantly impacted invasibility. Using path analysis, we found that community composition (dominant species identity) reduced invasion by reducing early-season light availability and increasing late-season plant community biomass. Nitrogen availability was an important factor for the survival of invaders in the second year of the experiment. We also found significant direct effects of certain dominant species on invasion, although the mechanisms driving these effects remain unclear. The magnitude of dominant species effects on invasibility we observed are comparable to species richness effects observed in other studies, showing that species composition and dominant species can have strong effects on the invasibility of a community.     

不同干扰强度对沙质海岸带植物物种β多样性的影响

选择东山岛沙质海岸带4种不同干扰强度林分作为空间演替系列,采用样方法研究其β多样性的变化,结果表明：（1）不同干扰强度林分的植物组成和重要值差别较大,在天然次生林的乔木层和灌木层中,潺槁木姜子（Litsea glutinosa）均是第一优势种,重要值分别为98和137,群落结构稳定;择伐干扰群落中,潺槁木姜子变为乔木层第三优势种,重要值为33,灌木层中仍为第一优势种,重要值为128;皆伐干扰群落中其优势地位逐渐降低,乔木层已无潺槁木姜子的分布,群落结构简单、稳定性差;（2）β多样性（CJ、CS、βWS、βC、βR、CN、CMH）测度表明,随着干扰强度的增大,群落间物种更替速率由低到高呈现增大趋势,物种变化较大,共有种逐渐减少;研究结果支持了＂初始植物区系＂学说,为选择合适树种构建稳定高效的沿海防护林体系提供了基础资料。     

人为干扰对江油地区马尾松人工林群落结构和物种多样性的影响

物种多样性是群落功能复杂性和稳定性的重要量度指标,人为干扰对森林群落的影响可以通过群落结构和物种多样性的变化而直接显现,不同干扰强度下的森林群落结构和物种多样性特征不同。采用典型样地法,对比研究两种人为干扰强度（干扰强和弱）对江油地区的马尾松（Pinus massoniana）人工林群落的物种组成、群落结构和物种多样性的影响,采用物种丰富度指数D值、Shannon-wienner指数H值、Simpson优势度指数H′值和均匀度指数Jsw值来综合衡量群落的物种多样性水平。结果表明,（1）在总面积3200 m2的8个样地中记录到124个物种,隶属于105属56科,干扰强的群落各层次物种数均低于干扰弱的群落。（2）不同强度人为干扰下马尾松人工林的群落结构不同,干扰弱的群落径级结构呈近正态分布,在径级Ⅵ（7 cm≤d〈9 cm）处出现数量的峰值26株,群落结构较稳定;干扰强的群落径级结构不呈正态分布,在径级Ⅶ（13 cm≤d〈15 cm）处出现峰值43株。干扰弱和干扰强的群落高度级结构与其径级结构具有相似性,干扰弱的群落的个体主要集中在低中高度级（3≤H〈17）;干扰强的群落的个体数在各高度级分布不均匀,主要分布在中高高度级（H≥13）;干扰强的群落的个体数在各径级和高度级分布相对不均匀,群落结构趋于不稳定。（3）人为干扰对马尾松人工林群落造成负面影响,随着人为干扰强度的增加,物种丰富度指数D值、Shannon-wiener多样性指数H值、均匀度多样性指数Jsw值降低,Simpson优势度指数H′值增大,物种多样性呈下降趋势。     

重庆库区不同海拔段繁殖鸟类群落的物种多样性

1996~1997年,对三峡工程重庆库区的鸟类群落的物种组成和种群数量进行了调查,共调查记录库区繁殖鸟类12目40科170种,数量17326只,其中留鸟有104种,占繁殖鸟总数的61.2%,夏侯鸟66种,占38.8%.用秩-多度曲线对不同海拔段的繁殖鸟类群落进行多样性分析的结果显示:1)秩-多度曲线方法的结果比Shannon-Wiener等物种多样性指数更直观,能更全面地反映多样性状况,该曲线可以对不同群落或相同群落在不同时间的物种多样性进行比较,是分析生物多样性的一种最直观的方法;2)人类干扰是影响各海拔段繁殖鸟类群落物种多样性的主要因素,人类干扰越强烈,鸟类群落物种多样性越低.图2表3参20     

Disturbance and resource availability act differently on the same suite of plant traits: revisiting assembly hypotheses

Understanding the mechanisms of trait selection at the scale of plant communities is a crucial step toward predicting community assembly. Although it is commonly assumed that disturbance and resource availability constrain separate suites of traits, representing the regenerative and established phases, respectively, a quantification and test of this accepted hypothesis is still lacking due to limitations of traditional statistical techniques. In this paper we quantify, using structural equation modeling (SEM), the relative contributions of disturbance and resource availability to the selection of suites of traits at the community scale. Our model specifies and reflects previously obtained ecological insights, taking disturbance and nutrient availability as central drivers affecting leaf, allometric, seed, and phenology traits in 156 (semi-) natural plant communities throughout The Netherlands. The common hypothesis positing that disturbance and resource availability each affect a set of mutually independent traits was not consistent with the data. Instead, our final model shows that most traits are strongly affected by both drivers. In addition, trait-trait constraints are more important in community assembly than environmental drivers in half of the cases. Both aspects of trait selection are crucial for correctly predicting ecosystem processes and community assembly, and they provide new insights into hitherto underappreciated ecological interactions.     

Priority effects and species sorting in a long paleoecological record of repeated community assembly through time

We studied the relative roles of environmental species sorting and priority effects in the assembly of ecological communities on long time scales, by analyzing community turnover of water fleas (Daphnia) in response to strong and recurrent environmental change in a fluctuating tropical lake. During the past 1800 years, Lake Naivasha (Kenya) repeatedly fluctuated between a small saline pond habitat during lowstands and a large freshwater lake habitat during highstands. Starting from a paleoecological reconstruction, we estimated the role of priority effects in Daphnia community assembly across 16 of these habitat turnovers and compared this with the response of the community to reconstructed changes in three environmental variables important for species sorting. Our results indicate that the best predictor of Daphnia community composition during highstands was the community composition just prior to the transition from lowstands to highstands. This reflects a long-lasting priority effect of late lowstand communities on highstand communities, arising when remnant lowstand populations fill newly available ecological space in the rapidly expanding lake habitat. Species sorting and priority effects had a comparable but relatively small influence on community composition during the lowstands. Moreover, these priority effects decayed rapidly with time as Daphnia communities responded to environmental change, in contrast with the highstand communities where priority effects lasted for several decades.     

The role of sponge competition on coral reef alternative steady states

Sponges constitute an abundant and functionally important component of coral reef systems. Given their demonstrated resistance to environmental stress, it might be expected that the role of sponges in reef systems under modern regimes of frequent and severe disturbance may become even more substantial. Disturbances have recently reshaped the community structure of many Caribbean coral reefs shifting them towards a state of persistent low coral cover and often a dominance of macroalgae. Using competition and growth rates recorded from Glover's Atoll in Belize, we parameterise a mathematical model used to simulate the three-way competition between sponges, macroalgae and coral. We use the model to determine the range of parameters in which each of the three species might be expected to dominate. Emergent properties arise from our simple model of this complex system, and these include a special case in which heightened competitive ability of macroalgae versus coral may counter-intuitively prove to be advantageous to the persistence of corals. Importantly, we show that even under scenarios whereby sponges fail to invade the system, inclusion of this third antagonist can qualitatively affect the likelihood of alternative stable states - generally in favour of macroalgal dominance. The interplay between multi-species competition and predation is complex, but our efforts highlight a key process that has, until now, remained unexplored: the extent to which sponges dissipate algal grazing pressure by providing generalist fish with an alternative food source. We highlight the necessity of identifying the extent by which this process takes place in tropical systems in order to improve projections of alternative stable states for Caribbean coral reefs.     

Type of disturbance and ecological history determine structural stability.

This study aims to reveal whether complexity, namely, community and trophic structure, of chronically stressed soil systems is at increased risk or remains stable when confronted with a subsequent disturbance. Therefore, we focused on a grassland with a history of four centuries of patchy contamination. Nematodes were used as model organisms because they are an abundant and trophically diverse group and representative of the soil food web and ecosystem complexity. In a field survey, a relationship between contaminants and community structures was established. Following, two groups of soil mesocosms from the field that differed in contamination level were exposed to different disturbance regimes, namely, to the contaminant zinc and a heat shock. The zinc treatment revealed that community structure is stable, irrespective of soil contamination levels. This implies that centuries of exposure to contamination led to adaptation of the soil nematode community irrespective of the patchy distribution of contaminants. In contrast, the heat shock had adverse effects on species richness in the highly contaminated soils only. The total nematode biomass was lower in the highly contaminated field samples; however, the biomass was not affected by zinc and heat treatments of the mesocosms. This means that density compensation occurred rapidly, i.e., tolerant species quickly replaced sensitive species. Our results support the hypothesis that the history of contamination and the type of disturbance determine the response of communities. Despite that ecosystems may be exposed for centuries to contamination and communities show adaptation, biodiversity in highly contaminated sites is at increased risk when exposed to a different disturbance regime. We discuss how the loss of higher trophic levels from the entire system, such as represented by carnivorous nematodes after the heat shock, accompanied by local biodiversity loss at highly contaminated sites, may result in detrimental effects on ecosystem functions.     

Effects of physical disturbance on nematode communities in sand and mud: a microcosm experiment

A microcosm experiment was carried out to evaluate the effects of continuous and spasmodic physical disturbance of differing frequency on the structure of nematode communities of intertidal sand and mud. There was a marked, characteristic change in abundance and diversity for both sediment types. In the sand microcosms, the majority of univariate measures of community structure, including species diversity, were lowest in the sediments subjected to a high frequency of disturbance. For the mud microcosms, most univariate measures reached their highest values in the treatments with an intermediate frequency of disturbance and were lower in treatments subjected to both higher and lower frequencies. Multivariate ordinations for both nematode assemblages showed a clear separation of undisturbed controls and disturbed treatments, but only for the muddy sediment was there a graded change in community composition with increasing frequency of disturbance. These results confirmed our a priori expectation that nematode assemblages from mobile sandy sediments would be more resilient to physical disturbance than those from sheltered muds, and these observations are considered in the context of Connell's intermediate disturbance hypothesis. Received: 7 May 1997 / Accepted: 10 October 1997     

Multidimensional trade-offs in species responses to disturbance: implications for diversity in a subtropical forest

Species employ diverse strategies to cope with natural disturbance, but the importance of these strategies for maintaining tree species diversity in forests has been debated. Mechanisms that have the potential to promote tree species coexistence in the context of repeated disturbance include life history trade-offs in colonization and competitive ability or in species' ability to survive at low resource conditions and exploit the temporary resource-rich conditions often generated in the wake of disturbance (successional niche). Quantifying these trade-offs requires long-term forest monitoring and modeling. We developed a hierarchical Bayes model to investigate the strategies tree species employ to withstand and recover from hurricane disturbance and the life history trade-offs that may facilitate species coexistence in forests subject to repeated hurricane disturbance. Unlike previous approaches, our model accommodates temporal variation in process error and observations from multiple sources. We parameterized the model using growth and mortality data from four censuses of a 16-ha plot taken every five years (1990-2005), together with damage data collected after two hurricanes and annual seed production data (1992-2005). Species' susceptibilities to hurricane damage as reflected by changes in diameter growth and fecundity immediately following a storm were weak, highly variable, and unpredictable using traditional life history groupings. The lower crowding conditions (e.g., high light) generated in the wake of storms, however, led to greater gains in growth and fecundity for pioneer and secondary-forest species than for shade-tolerant species, in accordance with expectation of life history. We found moderate trade-offs between survival in high crowding conditions, a metric of competitive ability, and long-distance colonization. We also uncovered a strong trade-off between mean species fecundity in low crowding conditions, a metric of recovery potential, and competitive ability. Trade-offs in competitive and colonization ability, in addition to successional niche processes, are likely to contribute to species persistence in these hurricane-impacted forests. The strategies species employ to cope with hurricane damage depend on the degree to which species rely on sprouting, repair of adult damage, changes in demographic rates in response to enhanced resource availability after storms, or long-distance dispersal as recovery mechanisms.     

Predicting Richness of Native, Rare, and Exotic Plants in Response to Habitat and Disturbance Variables across a Variegated Landscape

Species richness of native, rare native, and exotic understorey plants was recorded at 120 sites in temperate grassy vegetation in New South Wales. Linear models were used to predict the effects of environment and disturbance on the richness of each of these groups. Total native species and rare native species showed similar responses, with richness declining on sites of increasing natural fertility of parent material as well as declining under conditions of water enrichment (resulting from human-induced changes in drainage characteristics, leading to increased run-off), severe livestock grazing, and soil disturbance. The response of rare native species to water enrichment, however, was significantly greater than that of all native species. Exotic species richness varied in reverse to that of native species with positive responses to water enrichment and soil disturbance. The contrasting behaviors are attributed to differences in the evolutionary history of native and exotic assemblages and their resulting preadaptations to a landscape recently subjected to agricultural settlement. It would appear that for exogenous disturbances, the intermediate disturbance hypothesis is not supported by our data. In the sampled region, pastures represent the major land-use in terms of area, but have relatively low densities of native and rare species compared with more lightly grazed areas. However, their management is considered to be essential to the maintenance of diversity on a regional scale.