Resource pulses,species interactions,and diversity maintenance in arid and semi-arid environments

Arid environments are characterized by limited and variable rainfall that supplies resources in pulses. Resource pulsing is a special form of environmental variation, and the general theory of coexistence in variable environments suggests specific mechanisms by which rainfall variability might contribute to the maintenance of high species diversity in arid ecosystems. In this review, we discuss physiological, morphological, and life-history traits that facilitate plant survival and growth in strongly water-limited variable environments, outlining how species differences in these traits may promote diversity. Our analysis emphasizes that the variability of pulsed environments does not reduce the importance of species interactions in structuring communities, but instead provides axes of ecological differentiation between species that facilitate their coexistence. Pulses of rainfall also influence higher trophic levels and entire food webs. Better understanding of how rainfall affects the diversity, species composition, and dynamics of arid environments can contribute to solving environmental problems stemming from land use and global climate change.     

Moving from pattern to process: coexistence mechanisms under intermediate disturbance regimes

Coexistence mechanisms that require environmental variation to operate contribute importantly to the maintenance of biodiversity. One famous hypothesis of diversity maintenance under disturbance is the intermediate disturbance hypothesis (IDH). The IDH proposes patterns of peaked diversity under intermediate disturbance regimes, based on a tension between competitively superior species and species which can rapidly colonize following disturbance. We review the literature, and describe recent research that suggests that more than one underlying mechanism can generate this unimodal diversity pattern in disturbed environments. Several exciting emerging research areas are identified, including interactions between disturbance types, operation of the IDH in multi‐trophic systems, and changes in disturbance regimes. However, empirical work is still focussed on describing the IDH pattern, with little emphasis on identifying its mechanistic basis. We discuss how to extend methods for identifying different coexistence mechanisms, developed in the theoretical literature, to experimental research. In an attempt to operationalize these various ideas we outline a hypothetical IDH research programme. A solid understanding of the life history attributes of the component species and their responses to disturbance will facilitate identification of the coexistence mechanism(s) underlying the IDH pattern, and provide a framework by which empirical and theoretical results can be more fully integrated.     

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.     

Spatial patterns and coexistence mechanisms in systems with unidirectional flow

River ecosystems are the prime example of environments where unidirectional flow influences the dispersal of individuals. Spatial patterns of community composition and species replacement emerge from complex interplays of hydrological, geochemical, biological, and ecological factors. Local processes affecting algal dynamics are well understood, but a mechanistic basis for large scale emerging patterns is lacking. To understand how these patterns could emerge in rivers, we analyze a reaction-advection-diffusion model for two competitors in heterogeneous environments. The model supports waves that invade upstream up to a well-defined "upstream invasion limit". We discuss how these waves are produced and present their key properties. We suggest that patterns of species replacement and coexistence along spatial axes reflect stalled waves, produced from diffusion, advection, and species interactions. Emergent spatial scales are plausible given parameter estimates for periphyton. Our results apply to other systems with unidirectional flow such as prevailing winds or climate-change scenarios.     

Hutchinson revisited: Patterns of density regulation and the coexistence of strong competitors

Ecologists have long been searching for mechanisms of species coexistence, particularly since G.E. Hutchinson raised the ‘paradox of the plankton’. A promising approach to solve this paradox and to explain the coexistence of many species with strong niche overlap is to consider over-compensatory density regulation with its ability to generate endogenous population fluctuations.Previous work has analysed the role of over-compensation in coexistence based on analytical approaches. Using a spatially explicit time-discrete simulation model, we systematically explore the dynamics and conditions for coexistence of two species. We go beyond the analytically accessible range of models by studying the whole range of density regulation from under- to very strong over-compensation and consider the impact of spatial structure and temporal disturbances. In particular, we investigate how coexistence can emerge in different types of population growth models.We show that two strong competitors are able to coexist if at least one species exhibits over-compensation. Analysing the time series of population dynamics reveals how the differential responses to density fluctuations of the two competitors lead to coexistence: The over-compensator generates density fluctuations but is the inferior competitor at strong amplitudes of those fluctuations; the competitor, therefore, becomes frequent and dampens the over-compensator's amplitudes, but it becomes inferior under dampened fluctuations.These species interactions cause a dynamic alternation of community states with long-term persistence of both species. We show that a variety of population growth models is able to reproduce this coexistence although the particular parameter ranges differ among the models. Spatial structure influences the probability of coexistence but coexistence is maintained for a broad range of dispersal parameters.The flexibility and robustness of coexistence through over-compensation emphasize the importance of nonlinear density dependence for species interactions, and they also highlight the potential of applying more flexible models than the classical Lotka-Volterra equations in community ecology.     

Factors and mechanisms that explain coexistence in a Mediterranean carnivore assemblage: An integrated study based on camera trapping and diet

To promote management and conservation, it is useful to identify the factors that determine species distribution and to understand the mechanisms that regulate the organization of species assemblages or influence the dynamics of communities. Using information provided by 842 camera-trap photos and 8175 scats, we studied the factors that favour the coexistence of European badgers (Meles meles), red foxes (Vulpes vulpes) and stone martens (Martes foina) in a Mediterranean, agroforestry environment in the Iberian Peninsula. With extensive, simultaneous occupation of the space, and simultaneous activity during a broad time period (basically nocturnal and crepuscular activity patterns), the carnivores displayed different strategies depending on the availability of resources. In summer when plant resources were abundant and easy to access, there was a high overlap in patterns of diet and activity, and the temporal avoidance of the superior competitor allowed joint use of the same plots. In autumn, when there were fewer resources (although still sufficient) that were harder to access, the maintenance of food overlap was compensated for by avoidance and a reduction in overlapping activity. In winter and spring, the differentiation in response behaviour was evident in the partial substitution of plant resources.Differentiation in niche dimensions has been linked to complementarity, the differential needs and capacities of each species, and their biology. Differentiation in response behaviour was compatible with the hierarchical structure of the carnivores: European badger; red fox; stone marten. Knowledge of these factors and mechanisms increases our understanding and can help in the prediction of responses to disturbances. Consequently, it helps to improve management and conservation actions.     

Bioaugmentation and coexistence of two functionally similar bacterial strains in aerobic granules

The survival of the inoculated microbial culture is critical for successful bioaugmentation but impossible to predict precisely. As an alternative strategy, bioaugmentation of a group of microorganisms may improve reliability of bioaugmentation. This study evaluated simultaneous bioaugmentation of two functionally similar bacterial strains in aerobic granules. The two strains, Pandoraea sp. PG-01 and Rhodococcus erythropolis PG-03, showed high phenol degradation and growth rates in phenol medium, but they were characterized as having a poor aggregation activity and weak bioflocculant-producing and biofilm-forming abilities. In the spatially homogeneous batch conditions, strain PG-01 with higher growth rates outcompeted strain PG-03. However, the two strains could stably coexist in the spatially heterogeneous conditions. Then the two strains were mixed and bioaugmented into activated sludge in two sequencing batch reactors, which were operated with the different settling times of 5 and 30 min, respectively. Aerobic granules were developed only in the reactor with a settling time of 5 min. Fluorescence in situ hybridization and denaturing gradient gel electrophoresis showed that the two strains could coexist in aerobic granules but not in activated sludge. These findings suggested that the compact structure of aerobic granules provided spatial isolation for coexistence of competitively superior and inferior strains with similar functions.     

Variation in moisture duration as a driver of coexistence by the storage effect in desert annual plants

Temporal environmental variation is a leading hypothesis for the coexistence of desert annual plants. Environmental variation is hypothesized to cause species-specific patterns of variation in germination, which then generates the storage effect coexistence mechanism. However, it has never been shown how sufficient species differences in germination patterns for multispecies coexistence can arise from a shared fluctuating environment. Here we show that nonlinear germination responses to a single fluctuating physical environmental factor can lead to sufficient differences between species in germination pattern for the storage effect to yield coexistence of multiple species. We derive these nonlinear germination responses from experimental data on the effects of varying soil moisture duration. Although these nonlinearities lead to strong species asymmetries in germination patterns, the relative nonlinearity coexistence mechanism is minor compared with the storage effect. However, these asymmetries mean that the storage effect can be negative for some species, which then only persist in the face of interspecific competition through average fitness advantages. This work shows how a low dimensional physical environment can nevertheless stabilize multispecies coexistence when the species have different nonlinear responses to common conditions, as supported by our experimental data.     

Interacting coexistence mechanisms in annual plant communities: Frequency-dependent predation and the storage effect

We study frequency-dependent seed predation (FDP) in a model of competing annual plant species in a variable environment. The combination of a variable environment and competition leads to the storage-effect coexistence mechanism (SE), which is a leading hypothesis for coexistence of desert annual plants. However, seed predation in such systems demands attention to coexistence mechanisms associated with predation. FDP is one such mechanism, which promotes coexistence by shifting predation to more abundant plant species, facilitating the recovery of species perturbed to low density. When present together, FDP and SE interact, undermining each other’s effects. Predation weakens competition, and therefore weakens mechanisms associated with competition: here SE. However, the direct effect of FDP in promoting coexistence can compensate or more than compensate for this weakening of SE. On the other hand, the environmental variation necessary for SE weakens FDP. With high survival of dormant seeds, SE can be strong enough to compensate, or overcompensate, for the decline in FDP, provided predation is not too strong. Although FDP and SE may simultaneously contribute to coexistence, their combined effect is less than the sum of their separate effects, and is often less than the effect of the stronger mechanism when present alone.     

Scaling up from local competition to regional coexistence across two scales of spatial heterogeneity: insect larvae in the fruits of <Emphasis Type="Italic">Apeiba membranacea</Emphasis>

Species that live in patchy and ephemeral habitats can compete strongly for resources within patches at a small scale. The ramifications of these interactions for population dynamics and coexistence at regional scales will depend on the intraspecific and interspecific distributions of individuals among patches. Spatial heterogeneity due to independent aggregation of competitors among patchy habitats is an important mechanism maintaining species diversity. I describe regional patterns of aggregation for four species of insect larvae in the fruits of Apeiba membranacea, a Neotropical rainforest tree. This aggregation results from variation in densities at a small scale (among the fruits under a single tree), compounded by significant variation among trees in both mean densities and degrees of aggregation. Both the degrees of aggregation and mean densities are statistically independent within and across species at both spatial scales. I evaluate the regional consequences of these spatial patterns by using maximum likelihood methods to parameterize a model that includes both explicit measures of the strength of competition and spatial variation at both within- and among-tree spatial scales. Despite strong competitive interactions among these species, during 2 years the observed spatial variation at both scales combined was sufficient to explain the coexistence of these species, although other coexistence mechanisms may also operate simultaneously. The observed spatial variation at small spatial scales may not be sufficient for coexistence, indicating the importance of considering multiple sources of spatial heterogeneity when scaling up from experiments that investigate local interactions to regional patterns of coexistence.     

Quantifying the relative importance of variation in predation and the environment for species coexistence

Coexistence and food web theory are two cornerstones of the long‐standing effort to understand how species coexist. Although competition and predation are known to act simultaneously in communities, theory and empirical study of these processes continue to be developed largely independently. Here, we integrate modern coexistence theory and food web theory to simultaneously quantify the relative importance of predation and environmental fluctuations for species coexistence. We first examine coexistence in a theoretical, multitrophic model, adding complexity to the food web using machine learning approaches. We then apply our framework to a stochastic model of the rocky intertidal food web, partitioning empirical coexistence dynamics. We find the main effects of both environmental fluctuations and variation in predator abundances contribute substantially to species coexistence. Unexpectedly, their interaction tends to destabilise coexistence, leading to new insights about the role of bottom‐up vs. top‐down forces in both theory and the rocky intertidal ecosystem.     

Adaptive change in the resource-exploitation traits of a generalist consumer: the evolution and coexistence of generalists and specialists

Mathematical models of consumer-resource systems are used to explore the evolution of traits related to resource acquisition in a generalist consumer species that is capable of exploiting two resources. The analysis focuses on whether evolution of traits determining the capture rates of two resources by a consumer species produce one generalist, two specialists, or all three types, when all types are characterized by a common fitness function. In systems with a stable equilibrium, evolution produces one generalist or two specialists, depending on the second derivative of the trade-off relationship. When there are sustained population fluctuations, the nature of the trade-off between the consumer's capture rates of the two resources still plays a key role in determining the evolutionary outcome. If the trade-off is described by a choice variable between zero and one that is raised to a power n, polymorphic states are possible when n > 1, which implies a positive second derivative of the curve. These states are either dimorphism, with two relatively specialized consumer types, or trimorphism, with a single generalist type and two specialists. Both endogenously driven consumer-resource cycles, and fluctuations driven by an environmental variable affecting resource growth are considered. Trimorphic evolutionary outcomes are relatively common in the case of endogenous cycles. In contrast to a previous study, these trimorphisms can often evolve even when new lineages are constrained to have phenotypes very similar to existing lineages. Exogenous cycles driven by environmental variation in resource growth rates appear to be much less likely to produce a mixture of generalists and specialists than are endogenous consumer-resource cycles.     

Spatial and temporal patterns in the recruitment of the intertidal barnacle Chthamalus malayensis Pilsbry (Crustacea: Cirripedia) on the equatorial shores of Peninsular Malaysia and Singapore

Spatial and temporal variation in the recruitment of the intertidal barnacle Chthamalus malayensis was examined over one year (September 2003-August 2004) on the equatorial shores of southern Peninsular Malaysia and Singapore. A nested sampling design was applied for the first time on these shores, over three spatial scales - hundreds of kilometres, kilometres to tens of kilometres and tens of metres - and temporal variation was determined through monthly sampling of recruits. Shores within 2° north of the equator on the East and West coasts of southern Peninsular Malaysia and the southern coast of Singapore were selected as study sites. Generally, all three coasts recruited throughout the year, with varying intensities. There was a clear regional pattern, where the largest number of recruits was found on the East Coast of Malaysia, and the least in Singapore. Analyses of variance (ANOVA) showed that variability occurred on the smallest scale, but only in nine of the twelve months examined, consequently resulting in significant temporal and spatial interaction. Calculated variance components indicated that small-scale variation accounted for most of the overall variability. The potential causes of the spatial and temporal patterns of C. malayensis recruitment, and implications on tropical-temperate comparisons will be discussed.     

The coexistence and release of bovine pancreatic polypeptide-like immunoreactivity from noradrenergic superior cervical ganglia neurons

The coexistence of bovine pancreatic polypeptide-like (BPP) immunoreactivity within the cell bodies and axons of the superior cervical ganglia (SCG) was studied. Adjacent sections stained by the indirect immunofluorescence technique for either BPP or dopamine-β-hydroxylase (DBH) revealed that virtually all (90–95%) SCG cells contained DBH and a subpopulation (30–50%) also contained BPP. Ligation of the pre- and postganglionic nerves of the SCG demonstrated BPP-positive fibers emanating from the SCG via both axonal trunks. BPP-containing fibers were also observed entering the SCG via the preganglionic nerve. The peptide was co-released with catecholamine by electrical stimulation of the preganglionic cervical sympathetic nerve trunk, but was not effected by reserpine. The extensive distribution and unique coexistence of a PP-like peptide with peripheral sympathetic nerves suggests a neuromodulatory role in autonomic functions.     

克隆和有性亲体效应及其调控机制

植物表型受自身基因型、所处环境及其亲体所经历环境的共同影响;其中,亲体环境对子代表型的影响被称为亲体效应。亲体效应不仅可通过有性繁殖产生的种子传递给后代(即有性亲体效应),也可以通过克隆生长等无性繁殖产生的分株传递给后代(即克隆亲体效应)。亲体效应对植物种群,特别是对有性繁殖受限、缺乏遗传变异的克隆植物种群的长期进化可能发挥着极其重要的作用,因此,对亲体效应研究进展的梳理非常必要。对克隆亲体效应和有性亲体效应的内涵进行了阐释,并论述了克隆和有性亲体效应对子代表型、适合度、种内/种间竞争能力以及种群/群落结构和功能的潜在影响;阐述了亲体效应的潜在调控机制,包括供给机制、代谢物质调控机制、表观遗传机制等;论述了克隆亲体效应在克隆植物适应进化中的作用。未来可以就克隆亲体效应的遗传稳定性及其对克隆生活史性状变异的贡献程度,以及克隆和有性亲体效应引起的表型多样性对种内/种间关系、种群/群落多样性及生态系统结构、功能和稳定性的影响开展深入研究。     

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui Southward and Chthamalus stellatus (Poli) (Crustacea: Cirripedia) over an European scale

Spatial variation in the recruitment of the intertidal barnacles Chthamalus montagui and Chthamalus stellatus was examined over an European scale. The study was carried out using standardised protocols at a series of locations. The five locations chosen (SW Ireland, NW Spain, SW Portugal and NW and NE Italy) span a large part of the range of these species in Europe. The spatial scales were location (hundreds of kilometres) and shore (thousands of metres).Estimates of total cumulative recruitment (cyprids and metamorphs) summed over the year (April 1997 to March 1998) showed substantial variation between locations which was dependent on the species. Recruitment was highest in SW Portugal for C. montagui and in SW Ireland and NW Spain for C. stellatus. Overall recruitment of C. montagui was higher than that of C. stellatus at all locations except SW Ireland, where recruitment of the two species was not significantly different. There were significant differences among shores in each location.The recruitment period of both species varied with location, with recruitment beginning earlier further south. In general, recruitment of C. montagui and C. stellatus was recorded in 8 months in NW Spain and NE Italy, while only in 7 months in SW Ireland. Recruitment of C. montagui occurred in 10 months in SW Portugal, but no recruits of C. stellatus were found. In all locations there was at least one distinct peak of recruitment. In SW Ireland both species showed only one peak of recruitment, a month after initiation. At the more southerly Atlantic locations, as well as in the Mediterranean, two unequal peaks of recruitment were generally seen. During recruit census, the number of cyprids, in comparison to metamorphs, found at any location was very low. In SW Ireland and NW Spain cyprids of both species were found, while in SW Portugal and in the Mediterranean, only cyprids of C. montagui were found.     

Hamiltonian energy and coexistence of hidden firing patterns from bidirectional coupling between two different neurons

In this paper, bidirectional-coupled neurons through an asymmetric electrical synapse are investigated. These coupled neurons involve 2D Hindmarsh–Rose (HR) and 2D FitzHugh–Nagumo (FN) neurons. The equilibria of the coupled neurons model are investigated, and their stabilities have revealed that, for some values of the electrical synaptic weight, the model under consideration can display either self-excited or hidden firing patterns. In addition, the hidden coexistence of chaotic bursting with periodic spiking, chaotic spiking with period spiking, chaotic bursting with a resting pattern, and the coexistence of chaotic spiking with a resting pattern are also found for some sets of electrical synaptic coupling. For all the investigated phenomena, the Hamiltonian energy of the model is computed. It enables the estimation of the amount of energy released during the transition between the various electrical activities. Pspice simulations are carried out based on the analog circuit of the coupled neurons to support our numerical results. Finally, an STM32F407ZE microcontroller development board is exploited for the digital implementation of the proposed coupled neurons model.     

Patterns of adult abundance in Chthamalus stellatus (Poli) and C. montagui Southward (Crustacea: Cirripedia) emerge during late recruitment

The aim of this study was to investigate when adult distribution patterns are established in the barnacles Chthamalus stellatus and C. montagui. Adult ‘zones’ were identified by analysing field counts of both species at mid and upper shore heights. Monthly collections of cyprids, < 1 month old metamorphs and recruits (all metamorphosed individuals older than approximately 1 month) were made for C. stellatus and C. montagui in natural barnacle beds at six shores in SW Ireland. This was carried out over one year in 1996/1997, using a hierarchical sampling design. Abundance of total recruits (0-3 months old) was compared between adult zones after the main settlement season had ended. In addition, scales of variability in 0-3 month recruitment into adult zones were compared between the species at two scales: shores (1000s of metres) and sites within shores (10s of metres). Older recruits of each species, up to 11 months of age, were also compared between adult zones.The majority of settlement (measured as attached cyprids) occurred between August and October 1996. In October, there was no effect of adult zone on the abundance of total (0-3 month) recruitment up to that point in either species. Despite this homogeneity in recruitment between adult zones, significant spatial variation was found in 0-3 month recruits of both species at both of the scales examined. In C. stellatus the amount of variation associated with the larger scale (shore) was more than twice that of sites or of the residual variation (replicates within sites). 0-3 month recruitment in C. montagui was also most variable at the scale of shores but the residual variability (between replicates within site) was of similar magnitude to that of shores. Variability in 0-3 month C. montagui recruitment was relatively low at the scale of sites.There was a small but consistent input of recruits to adult zones over 9 months of the year, complicating the assessment of when adult patterns were set-up in these species. By June 1997, characteristic patterns of adult dominance had been established at all shores. Settlement had completely ceased by this time and individual barnacles were potentially 11 months old. Neither settlement nor early recruitment are significant in determining adult zonation patterns in these species. Instead, differential mortality patterns in individuals up to the age of 11 months are implicated in determining patterns of distribution of both species.     

On the mechanisms of coexistence among annual-plant species, using neighbourhood techniques and simulation models

Many studies have investigated the density-dependent regulation ofannual-plant populations on coastal sand dunes, but few have explored theconsequences of competition for the coexistence of plants in these simplecommunities. We used neighbourhood techniques to parameterize competition anddispersal functions from field data collected for two species of dune annual(Aira praecox and Erodium cicutarium)over three successive years, and then combined these functions into spatiallyexplicit simulation models. The population size of Airavaried enormously between years, while Erodium remainedsteady. Competition with neighbours reduced the spike length ofAira plants only in one of the three years (when itspopulation density was highest), while competition with neighbouringErodium plants appeared to result in the local death ofAira plants. However, these density-dependent effects werefar too weak to generate the observed changes in the population size ofAira among years, or to maintain populations below theupper limits observed. The large-seeded Erodium wasaffected by intraspecific competition but was unaffected by small-seededAira plants. Therefore, the larger-seeded species wascompetitively superior to the smaller-seeded species, an affect that couldpromote coexistence (albeit weakly) by a competition-colonisation trade-off.Modal dispersion distances of Aira andErodium were 45 and 60 mm respectively,greater than the radius within which competitive interactions occurred (40mm). Theoretical studies suggest that under these conditions thespatial arrangement of plants should be nearly random. In factAira was spatially aggregated, especially when rare,suggesting that patchy mortality across the dunes was important in generatingspatial structure. The study suggests that density dependence only weaklyregulates dune annual communities, while year-to-year environmental variationexert major influences on population sizes and spatial structures.     

The role of seasonal timing and phenological shifts for species coexistence

Shifts in the phenologies of coexistence species are altering the temporal structure of natural communities worldwide. However, predicting how these changes affect the structure and long‐term dynamics of natural communities is challenging because phenology and coexistence theory have largely proceeded independently. Here, I propose a conceptual framework that incorporates seasonal timing of species interactions into a well‐studied competition model to examine how changes in phenologies influence long‐term dynamics of natural communities. Using this framework I demonstrate that persistence and coexistence conditions strongly depend on the difference in species’ mean phenologies and how this difference varies across years. Consequently, shifts in mean and interannual variation in relative phenologies of species can fundamentally alter the outcome of interactions and the potential for persistence and coexistence of competing species. These effects can be predicted by how per‐capita effects scale with differences in species’ phenologies. I outline how this approach can be parameterized with empirical systems and discuss how it fits within the context of current coexistence theory. Overall, this synthesis reveals that phenology of species interactions can play a crucial yet currently understudied role in driving coexistence and biodiversity patterns in natural systems and determine how species will respond to future climate change.