Indirect interactions among tropical tree species through shared rodent seed predators: a novel mechanism of tree species coexistence

The coexistence of numerous tree species in tropical forests is commonly explained by negative dependence of recruitment on the conspecific seed and tree density due to specialist natural enemies that attack seeds and seedlings (‘Janzen–Connell’ effects). Less known is whether guilds of shared seed predators can induce a negative dependence of recruitment on the density of different species of the same plant functional group. We studied 54 plots in tropical forest on Barro Colorado Island, Panama, with contrasting mature tree densities of three coexisting large seeded tree species with shared seed predators. Levels of seed predation were far better explained by incorporating seed densities of all three focal species than by conspecific seed density alone. Both positive and negative density dependencies were observed for different species combinations. Thus, indirect interactions via shared seed predators can either promote or reduce the coexistence of different plant functional groups in tropical forest.     

The importance of temporal climate variability for spatial patterns in plant diversity

Spatial variation in absolute climatic conditions (means, maxima or minima) is widely acknowledged to play a fundamental role in controlling species diversity patterns. In contrast, while evidence is accumulating that variability around mean climatic conditions may also influence species coexistence and persistence, the importance of spatial variation in temporal climatic variability for species diversity is still largely unknown. We used a unique dataset capturing fine‐scale spatial heterogeneity in temperature variability across 2490 plots in southeast Australia to examine the comparative strength of absolute temperature and temperature variability in explaining spatial variation in plant diversity. Across all plots combined and in three of five forest types, temperature variability emerged as the better predictor of diversity. In all but one forest type, diversity also exhibited either a significant unimodal or positive linear correlation with temperature variability. This relationship is consistent with theory that predicts diversity will initially increase along a climate variability gradient due to temporal niche partitioning, but at an intermediary point, may decline as the risk of stochastic extinction exceeds competitive stabilization. These findings provide critical empirical evidence of a linkage between spatial variation in temporal climate variability and plant species diversity, and in light of changing climate variability regimes, highlight the need for ecologists to expand their purview beyond absolutes and averages.     

Concordant population dynamics of Lepidoptera herbivores in a forest ecosystem

It is frequently assumed that population fluctuations are largely independent within a community of trophically‐similar species, but this need not be so. If population fluctuations are partly synchronized or concordant, this will produce interannual variability in the community's aggregate abundance and generate temporal variance in ecosystem structure. We studied the community of Lepidoptera inhabiting northern hardwood forests in New Hampshire, USA, to evaluate the hypothesis that fluctuations in consumer communities can arise from concordant dynamics of constituent populations. Interannual comparisons of moth abundances for >75 species sampled at three sites over four years revealed that concordant dynamics contribute strongly to interannual variability in the abundance of consumers. A conspicuous decline in community abundance from 2004 to 2005 was the result of predominantly negative population growth rates of the component species, while an increase in community abundance from 2006 to 2007 was the result of predominantly positive population growth rates. Population dynamics most strongly linked species that feed in the early season (perhaps due to shared responses to climatic effects), but not species that might share natural enemies or host plants. The observed concordant dynamics introduced conspicuous temporal variation in the abundance of primary consumers relative to plants and secondary consumers, thereby altering the forest's trophic structure. Such variance in the aggregate abundance of forest primary consumers could generate time‐lagged fluctuations in abundances of secondary consumers and will generally have important consequences for ecosystem properties and processes that are nonlinear functions of consumer abundance, such as plant community structure and nutrient cycling.     

Tree diversity mitigates defoliation after a drought‐induced tipping point

Understanding the processes that underlie drought‐related tree vitality loss is essential for anticipating future forest dynamics, and for developing management plans aiming at increasing the resilience of forests to climate change. Forest vitality has been continuously monitored in Europe since the acid rain alert in the 1980s, and the intensive monitoring plots of ICP Forests offer the opportunity to investigate the effects of air pollution and climate change on forest condition. By making use of over 100 long‐term monitoring plots, where crown defoliation has been assessed extensively since 1990, we discovered a progressive shift from a negative to a positive effect of species richness on forest health. The observed tipping point in the balance of net interactions, from competition to facilitation, has never been reported from real ecosystems outside experimental conditions; and the strong temporal consistency of our observations with increasing drought stress emphasizes its climate change relevance. Furthermore, we show that higher species diversity has reduced the severity of defoliation in the long term. Our results confirm the greater resilience of diverse forests to future climate change‐induced stress. More generally, they add to an accumulating body of evidence on the large potential of tree species mixtures to face manifold disturbances in a changing world.     

Temporal niche dynamics, relative abundance and phylogenetic signal of coexisting species

The coexistence of similar species accounts for some 30% of diversity within communities, yet the coexistence and relative abundance of similar species is a continuing ecological conundrum. Using close phylogenetic relatedness as a measure of similarity, we previously demonstrated that neither classic niche theory nor neutral theory can explain the relative abundances of co-occurring pairs of similar tree species in a diverse tropical forest. Here, we show that the stable, focused competition of a temporal niche dynamic fits the distribution of observed fractional abundances (pairwise relative abundances). Previously published, independent evidence of temporal dynamics in this community supports our results; our model identifies additional criteria for field tests of differential sensitivity (DS) temporal dynamics. The success of temporal dynamics at explaining the observed distribution—and the failure of alternative hypotheses to do so—indicates that current diagnostics of community structure and assembly needs general re-examination. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of wildfires and post-fire forest treatments on rabbit abundance

One of the main factors involved in the decline in the European wild rabbit in the Iberian Peninsula is the loss of suitable habitats caused by abandonment of agricultural and grazing activities. Nowadays, Mediterranean landscapes suffer from wildfires that affect extensive areas and produce considerable habitat changes. However, little is known about the influence of wildfires and post-fire treatments on rabbit abundance to address policies to recover their populations. To do so, we studied abundances of this species in four types of plots during three consecutive years after a wildfire in Catalonia (NE Spain): (A) unburnt forests, (B) burnt forests with removal of burnt trees but with branches left, (C) burnt forests with removal of burnt trees and branches, and (D) non-forested burnt plots. Rabbits progressively colonised burnt plots, where their abundance increased for at least 5 years after the fire, but decreased or even disappeared in unburnt ones, indicating that forest fires have a positive effect on rabbit populations. Although abundances did not differ between the three burnt plot types, plots with removal of burnt branches had the highest increase in abundance. In addition, soil covered by branches or by dense vegetation appeared negatively correlated with abundance, indicating that this could hinder rabbit movements, while some plant species could benefit rabbits by providing high quality food. Thus, post-fire treatments favourable to rabbit populations might therefore be a good way of increasing the conservation and economic value of areas affected by forest fires.     

Niche partitioning and niche filtering jointly mediate the coexistence of three closely related spider species (Araneae,Philodromidae)

1. Several mechanisms can mediate the coexistence of species, such as the neutral dynamic, niche filtering and niche partitioning. The present study investigated which of these mechanisms mediate the coexistence of closely related spider species at the scale of one locality. 2. The coexistence of three spider species of the genus Philodromus (Philodromidae) (Philodromus albidus, Philodromus aureolus and Philodromus cespitum) was studied. The study area comprised three habitat types, including a deciduous forest, a scrub and a plum tree stand. The spatial niche properties of the Philodromus species were explored by comparing their micro/macrohabitat preferences. The natural diet of these species was analysed. Laboratory experiments involving prey acceptance were conducted to investigate the trophic niche properties. The species' phenologies were studied to compare their temporal niches. 3. The philodromids had differentiated their trophic and habitat niches. The coexistence mechanisms were therefore assessed by studying the relationships between niche overlaps and the pairwise product of relative abundances. A negative relationship was observed between functional niche overlaps and the product of relative abundances, whereas a positive relationship was observed between spatial niche overlaps and the product of relative abundances. 4. The present results suggest that different mechanisms influence different niche dimensions. Niche partitioning influenced functional niches, whereas niche filtering influenced spatial niches. The results also suggest that the filtering process in one dimension was facilitated by niche differentiation in another dimension.     

Dynamics of seedling populations and tree species coexistence in a forest: a simulation study

Theoretically, temporal variation of reproduction promotes species coexistence of sessile and polycarpic organisms when the reproduction is synchronized within species but independent among species. Monopoly of vacant sites and high relative population growth rate of minor species in the absence of propagules of other species is the essence of this mechanism. The mechanism is expected to work in forests, but persistent populations of seedlings may affect the promotion of coexistence. Using a tree-based simulation model of forest dynamics, it was demonstrated that the number of coexisting tree species was sensitively affected by the seedling establishment rate. The coexistence was not enhanced by temporal variation of reproduction when seedling establishment rate was low. This is because the reproducing minor species fail to monopolize vacant sites and allow the establishment of seedlings of other species in later years. High mortality of established seedlings under shade also suppressed coexistence. This is likely to be the result of a reduced storage effect of the population of seedlings. A forest structure and dynamics pattern that appears when tree species coexistence is promoted by fluctuating reproduction was searched for, and the number of coexisting species was varied by manipulating the seedling establishment rate. No distinct difference other than the species number itself was found between species-rich and species-poor forests. For example, the seedling population size varied, reflecting the temporal variation of reproduction, irrespective of the seedling establishment rate. Further strategy development is needed to validate the proposed mechanisms of species coexistence.     

Soil heterogeneity in tree mixtures depends on spatial clustering of tree species

Heterogeneity in soil characteristics promotes and maintains coexistence between a diverse set of species. In forests, trees have species-specific impacts on soil abiotic characteristics and mixing of tree species is being promoted as a tool to ensure high levels of diversity and functioning. Yet, limited knowledge is available on the effect of tree species composition and spatial clustering on heterogeneity in soil characteristics. In this paper we derived heterogeneity of key characteristics of the leaf litterfall, the forest floor and the mineral topsoil (C, N and base cation concentration, C:N ratio and mass) in 53 plots of 7 different tree species compositions. We found that heterogeneity increased from the leaf litterfall, through the forest floor down to the mineral topsoil. Mixing tree species did not lead to an increased heterogeneity in the forest floor and topsoil compared to monocultures. However, we did find that mixed plots where conspecific trees stand in groups are more heterogeneous than plots where species are intimately mixed. Our results imply that heterogeneity in soil characteristics does not necessarily increase with tree diversity, but that within mixed stands the spatial organization of tree species should be considered in relation to the scale at which heterogeneity is desired.     

Declining tree growth resilience mediates subsequent forest mortality in the US Mountain West

Climate change-triggered forest die-off is an increasing threat to global forests and carbon sequestration but remains extremely challenging to predict. Tree growth resilience metrics have been proposed as measurable proxies of tree susceptibility to mortality. However, it remains unclear whether tree growth resilience can improve predictions of stand-level mortality. Here, we use an extensive tree-ring dataset collected at ~3000 permanent forest inventory plots, spanning 13 dominant species across the US Mountain West, where forests have experienced strong drought and extensive die-off has been observed in the past two decades, to test the hypothesis that tree growth resilience to drought can explain and improve predictions of observed stand-level mortality. We found substantial increases in growth variability and temporal autocorrelation as well declining drought resistance and resilience for a number of species over the second half of the 20th century. Declining resilience and low tree growth were strongly associated with cross- and within-species patterns of mortality. Resilience metrics had similar explicative power compared to climate and stand structure, but the covariance structure among predictors implied that the effect of tree resilience on mortality could partially be explained by stand and climate variables. We conclude that tree growth resilience offers highly valuable insights on tree physiology by integrating the effect of stressors on forest mortality but may have only moderate potential to improve large-scale projections of forest die-off under climate change.     

Changes in forest bird abundance,community structure and composition following a hurricane in Sweden

The effect that extreme natural events have on biological diversity is relatively poorly known. We used a before–after control‐impact (BACI) design to analyze changes in bird abundances and communities following Hurricane Gudrun, which struck southern Sweden in January 2005, felling 75 million m³ of forest and causing damage to 5% of forested areas (half a million hectares) in a few hours. We used recent measures of impact in combination with classical BACI contrasts to analyze bird count data from a monitoring program in Sweden. We investigated changes in the abundance of 17 species commonly found in forests, along with changes in species composition and functional structure of the bird community. In total, we considered 34 response variables and examined whether responses were immediate or long‐term. There was no evidence of a strong effect of the hurricane on the abundances of six species. Estimates of the effects on five species were too uncertain to draw inferences. We detected positive and negative effects of the hurricane on the abundances of the remaining six species, but the magnitude of effects often was small. Generally, the effects were in the expected direction: negative on birds associated with mature forest and positive on birds associated with open land or young forest. We found evidence of an increase in the proportion of species that nest on the ground and a decrease in the proportion of species that nest in cavities and trees. In contrast, the hurricane had no discernible effect on functional measures of diversity (richness, evenness or divergence), or on communities’ reproductive or morphological characteristics. Our results suggest that the hurricane affected bird populations and communities, but the magnitude of effects was generally small.     

Understanding landscape patterns of temporal variability in avian populations to improve environmental impact assessments

It is recognized that wildlife populations exhibit spatial and temporal variability in patterns of species richness across heterogeneous landscapes. This phenomenon can prove problematic for environmental practitioners when attempting to complete comprehensive environmental assessments (EAs) with limited field surveys. A better understanding of regional spatio-temporal patterns in population dynamics should enhance site-level decision-making. In this study, the variability of seasonal data across the Credit River Watershed, southern Ontario, is assessed for a hierarchy of conservation measures including species richness, and two conservation wildlife response guilds based on primary habitat and area sensitivity. Bird populations were monitored at 24 forest monitoring plots across the watershed by the authors twice a season from 2003 to 2010 following the protocol of Environment Canada's Forest Bird Monitoring Program. The monitoring plots are located within four land management zones identified as 1) urban, 2) transitional, 3) escarpment and 4) rural. Data from the monitoring program are used to compare species richness among plots across the watershed and among land management zones. In addition, the variability of records from each plot over the 8 year period was determined by means of the Coefficient of Variation (CV) statistic. The mean variability of these records at each site within each land management zone was determined in order to assess whether the temporal variability of bird records might affect the integrity of short term assessments. Finally, a one-way ANOVA was applied to learn whether the result of short-term assessments may be further compounded by differences in the response of selected bird guilds to landscape heterogeneity. The results show that there is a significant difference in mean richness of forest birds among the four management zones. The ANOVAs indicate that significant difference is due to the temporal variability of a) breeding forest interior birds rather than edge birds or generalist species and b) breeding area sensitive species rather than area non-sensitive species. Recommendations are made that environmental assessments targeting forest interior bird populations need to plan sampling strategies that recognize this variability, especially for sites within the transitional and urban zones. Planning in the transition or urbanizing landscape should incorporate landscape ecology principles to sustain current levels of richness in forest species.     

The relationship between species richness and ecosystem variability is shaped by the mechanism of coexistence

Theory relating species richness to ecosystem variability typically ignores the potential for environmental variability to promote species coexistence. Failure to account for fluctuation‐dependent coexistence may explain deviations from the expected negative diversity–ecosystem variability relationship, and limits our ability to predict the consequences of increases in environmental variability. We use a consumer‐resource model to explore how coexistence via the temporal storage effect and relative nonlinearity affects ecosystem variability. We show that a positive, rather than negative, diversity–ecosystem variability relationship is possible when ecosystem function is sampled across a natural gradient in environmental variability and diversity. We also show how fluctuation‐dependent coexistence can buffer ecosystem functioning against increasing environmental variability by promoting species richness and portfolio effects. Our work provides a general explanation for variation in observed diversity–ecosystem variability relationships and highlights the importance of conserving regional species pools to help buffer ecosystems against predicted increases in environmental variability.     

Shade tolerance,canopy gaps and mechanisms of coexistence of forest trees

The belief that canopy gaps are important for the maintenance of tree species diversity appears to be widespread, but there have been no formal theoretical models to assess under what conditions gap phase processes allow coexistence. Much of the empirical research on niche differentiation in response to gaps has focused on evidence for an interspecific tradeoff between low light survival and high light growth. The objectives of this study are first to distinguish the possible mechanisms allowing coexistence based on this tradeoff, and second, to explore their limitations. We present a theory of forest dynamics driven by small‐scale disturbances as a special case of the theory of coexistence in variable environments. We demonstrate that temporal and spatial heterogeneity in light conditions that results from canopy gaps can allow stable coexistence as a result of three previously documented general mechanisms: ‘relative non‐linearity’, ‘the successional niche’ and the ‘storage effect’. We find that temporal fluctuations in light availability alone allow the stable coexistence of only two species. Spatial variation in disturbance synchronicity and intensity allows three species to coexist in a narrow parameter space. The rate of extinction is, however, extremely slow and there is transient coexistence of a larger number of species for a long period of time. We conclude that while the low light survival/high light growth tradeoff may be ubiquitous in forest tree species, it is unlikely to function as an important mechanism for the stable coexistence of several tree species.     

How do trees respond to species mixing in experimental compared to observational studies?

For decades, ecologists have investigated the effects of tree species diversity on tree productivity at different scales and with different approaches ranging from observational to experimental study designs. Using data from five European national forest inventories (16,773 plots), six tree species diversity experiments (584 plots), and six networks of comparative plots (169 plots), we tested whether tree species growth responses to species mixing are consistent and therefore transferrable between those different research approaches. Our results confirm the general positive effect of tree species mixing on species growth (16% on average) but we found no consistency in species‐specific responses to mixing between any of the three approaches, even after restricting comparisons to only those plots that shared similar mixtures compositions and forest types. These findings highlight the necessity to consider results from different research approaches when selecting species mixtures that should maximize positive forest biodiversity and functioning relationships.     

Structural and Floristic Heterogeneity in a 30-Year-Old Costa Rican Rain Forest Restored on Pasture Through Natural Secondary Succession

Relatively little information exists on neotropical secondary rain forests that have progressed beyond the pioneer stages of succession, or on the potential of natural regeneration to restore forest on large areas. We determined the structural and floristic characteristics (10 cm dbh) of a 30‐year‐old secondary forest developing on a 32.5 ha pasture on hilly terrain, abandoned after use of moderate intensity. Ten 0.24 ha sample plots covered the range of site conditions. The forest was dominated by long‐lived pioneer tree species; overall, the majority of species (70%) was vertebrate dispersed but the majority of individuals (52%) was of wind‐dispersed species. Tree species, including the dominants, were a mixture of those present in old‐growth and adventives colonizing from agricultural land. The forest was very heterogeneous. Vochysia ferruginea‐dominated stands characterized slopes with soils of high exchangeable acidity, while the adventive Cordia alliodora dominated sites with gentler topography and soils of lower acidity. Structural differences between the two forest types were slight, but Cordia forest had significantly greater species diversity and absolute and relative abundances of vertebrate‐dispersed tree species than Vochysia forest, which had significantly greater absolute and relative abundances of wind‐dispersed tree species. These latter differences between forest types, as well as the wide structural variation of the forest as a whole, were probably largely due to spatial and temporal variation in seed rains, some of it linked to the characteristics of the dominant species. Rain forest restoration on large pastures may depend greatly on wind dispersal and adventive tree species, and techniques for silvicultural diagnosis must be developed as a basis for the management of heterogeneous successional stands. Studies of early colonization of pastures should be expanded to focus on the causes of heterogeneity in older forests.     

Do lianas shape ant communities in an early successional tropical forest?

Almost half of lowland tropical forests are at various stages of regeneration following deforestation or fragmentation. Changes in tree communities along successional gradients have predictable bottom‐up effects on consumers. Liana (woody vine) assemblages also change with succession, but their effects on animal succession remain unexplored. Here we used a large‐scale liana removal experiment across a forest successional chronosequence (7–31 years) to determine the importance of lianas to ant community structure. We conducted 1,088 surveys of ants foraging on and living in trees using tree trunk baiting and hand‐collecting techniques at 34 paired forest plots, half of which had all lianas removed. Ant species composition, β‐diversity, and species richness were not affected by liana removal; however, ant species co‐occurrence (the coexistence of two or more species in a single tree) was more frequent in control plots, where lianas were present, versus removal plots. Forest stand age had a larger effect on ant community structure than the presence of lianas. Mean ant species richness in a forest plot increased by ca. 10% with increasing forest age across the 31‐year chronosequence. Ant surveys from forest >20 years old included more canopy specialists and fewer ground‐nesting ant species versus those from forests <20 years old. Consequently, lianas had a minimal effect on arboreal ant communities in this early successional forest, where rapidly changing tree community structure was more important to ant species richness and composition.     

Changes in tree reproductive traits reduce functional diversity in a fragmented Atlantic forest landscape

Functional diversity has been postulated to be critical for the maintenance of ecosystem functioning, but the way it can be disrupted by human-related disturbances remains poorly investigated. Here we test the hypothesis that habitat fragmentation changes the relative contribution of tree species within categories of reproductive traits (frequency of traits) and reduces the functional diversity of tree assemblages. The study was carried out in an old and severely fragmented landscape of the Brazilian Atlantic forest. We used published information and field observations to obtain the frequency of tree species and individuals within 50 categories of reproductive traits (distributed in four major classes: pollination systems, floral biology, sexual systems, and reproductive systems) in 10 fragments and 10 tracts of forest interior (control plots). As hypothesized, populations in fragments and control plots differed substantially in the representation of the four major classes of reproductive traits (more than 50% of the categories investigated). The most conspicuous differences were the lack of three pollination systems in fragments--pollination by birds, flies and non-flying mammals--and that fragments had a higher frequency of both species and individuals pollinated by generalist vectors. Hermaphroditic species predominate in both habitats, although their relative abundances were higher in fragments. On the contrary, self-incompatible species were underrepresented in fragments. Moreover, fragments showed lower functional diversity (H' scores) for pollination systems (-30.3%), floral types (-23.6%), and floral sizes (-20.8%) in comparison to control plots. In contrast to the overwhelming effect of fragmentation, patch and landscape metrics such as patch size and forest cover played a minor role on the frequency of traits. Our results suggest that habitat fragmentation promotes a marked shift in the relative abundance of tree reproductive traits and greatly reduces the functional diversity of tree assemblages in fragmented landscapes.     

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.     

Variability in solar radiation and temperature explains observed patterns and trends in tree growth rates across four tropical forests

The response of tropical forests to global climate variability and change remains poorly understood. Results from long-term studies of permanent forest plots have reported different, and in some cases opposing trends in tropical forest dynamics. In this study, we examined changes in tree growth rates at four long-term permanent tropical forest research plots in relation to variation in solar radiation, temperature and precipitation. Temporal variation in the stand-level growth rates measured at five-year intervals was found to be positively correlated with variation in incoming solar radiation and negatively related to temporal variation in night-time temperatures. Taken alone, neither solar radiation variability nor the effects of night-time temperatures can account for the observed temporal variation in tree growth rates across sites, but when considered together, these two climate variables account for most of the observed temporal variability in tree growth rates. Further analysis indicates that the stand-level response is primarily driven by the responses of smaller-sized trees (less than 20 cm in diameter). The combined temperature and radiation responses identified in this study provide a potential explanation for the conflicting patterns in tree growth rates found in previous studies.