Quantifying the relative importance of natural variables,human disturbance and spatial processes in ecological status indicators of boreal lakes

Spatial processes are increasingly associated with species distributions in freshwaters. However, these processes are usually neglected in bioassessment techniques, which may introduce uncontrolled variation in ecological indicators used to express human disturbance. We used partial linear regression to quantify the relative importance of natural variables, human disturbance and spatial variables in structuring variation in boreal lake status indicators based on six biological indicator groups (phytoplankton, macrophytes, diatoms, littoral and profundal macroinvertebrates and fish). We found that, of the pure fractions, human disturbance explained most variation (7–32%) of the ecological quality ratios (EQRs) for all groups, with the exception of littoral macroinvertebrate metric, which was most controlled by natural and spatial variables (15% and 16%, respectively). In addition, pure fractions of natural and spatial variables and joint fractions of different explanatory variable groups structured all biological metrics to various degrees. Phytoplankton, diatom and profundal macroinvertebrate EQRs responded purest to human disturbance but only weakly to pure natural or spatial variation. Our work demonstrates that spatial processes and spatial structuring of the environment can bias bioassessment techniques and hinder the detection of human impact. Thus, it is important to acknowledge spatial autocorrelation, context of metacommunity dynamics, species dispersal traits and variable spatial extent when defining reference conditions and bioassessment techniques for different biological groups. More research is needed to better understand the relative role of spatial processes on ecological metrics originated from different freshwater ecosystems. To this end, our work provides an example of how sources of variation can be identified to increase accuracy in freshwater bioassessment.     

Demographic analysis of Hubbell's neutral theory of biodiversity

Hubbell's neutral model is increasingly applied in both theoretical and empirical studies but so far little attention has been paid to the ecological mechanisms that determine species diversity in neutral communities. In this contribution we use a stochastic individual-based Markovian model to provide an explicit derivation of Hubbell's local community model from the fundamental processes of reproduction, mortality, and immigration, and show that such derivation provides important insights on the mechanisms regulating species diversity that cannot be obtained from the original model and its previous extensions. One important insight is that the basic parameters of Hubbell's model, community size (J) and the probability that a dying individual will be replaced by an immigrant (m), cannot be considered independent and that their interdependency leads to a counterintuitive trade-off between community size and species diversity. We further demonstrate that Hubbell's treatment of community size as a free parameter hides fundamental mechanisms that influence species diversity through their effect on the size of the community. For example, while in Hubbell's model immigration can only increase species diversity by promoting colonization rates, the demographic derivation shows that immigration can also promote species diversity by reducing extinction rates. Our demographic derivation also unifies previous contrasting predictions about the effect of reproduction on species diversity by showing that both positive and negative effects are possible, and that the balance between the two effects depends on the size of the community. The demographic derivation also reconciles an apparent contradiction between Hubbell's theory and patch occupancy theory, and integrates three previously proposed mechanisms of species diversity, the More Individuals Hypothesis, the rescue effect, and the dilution effect, within a single, unified framework.     

Neutral Models with Generalised Speciation

Hubbell’s neutral theory claims that ecological patterns such as species abundance distributions can be explained by a stochastic model based on simple assumptions. One of these assumptions, the point mutation assumption, states that every individual has the same probability to speciate. Etienne et al. have argued that other assumptions on the speciation process could be more realistic, for example, that every species has the same probability to speciate (Etienne, et al. in Oikos 116:241–258, 2007). They introduced a number of neutral community models with a different speciation process, and conjectured formulas for their stationary species abundance distribution. Here we study a generalised neutral community model, encompassing these modified models, and derive its stationary distribution, thus proving the conjectured formulas.     

Topographic and spatial controls of palm species distributions in a montane rain forest, southern Ecuador

The northern Andes harbour a flora that is as species-rich or even richer than the 18-times larger lowland Amazon basin. Gaining an understanding of how the high species richness of the Andean region is generated and maintained is therefore of particular interest. Environmental sorting due to elevational gradients in climate has been emphasized as a driver of vegetation distribution and plant community assembly in tropical mountain areas such as the Andes for two centuries, while alternative mechanisms have been little studied. Here, we investigated the importance of topography and spatial location as factors controlling species distributions in a palm community in a montane rain forest landscape in the Andes of southern Ecuador (1900–2150 m above sea level). Eleven species were present: Aiphanes verrucosa, Ceroxylon parvifrons, Chamaedorea pinnatifrons, Dictyocaryum lamarckianum, Euterpe precatoria, Geonoma densa, Geonoma orbignyana, Geonoma paradoxa, Prestoea acuminata and Wettinia aequatorialis. To study their spatial distribution, forty 250 m² (5 × 50 m²) plots were laid out perpendicular to four paths that were categorized into three areas and two topographic units (ridges and gullies). Mantel tests and indicator species analysis showed that both topography and spatial location imposed strong controls on palm species distributions at the study site. Our results suggest that species distributions in the studied montane forest landscape were partly determined by the species’ habitat requirements, but also by unknown spatial effects. Although a number of possible explanations exist for the latter, such as unmeasured environmental variables and historical disturbance events, we believe dispersal limitation is likely to be involved. Furthermore, although the gully- or ridge-association of some species corresponded to their general elevational ranges in southern Ecuador, this was not the case for other species. Based on such considerations, we conclude that elevational climatic gradients are likely to only form part of the explanation for the topographic effects on palm species distributions at the study site. Other factors must also be involved, notably wind-exposure and hydrology, as discussed for lowland palm communities. Our results show that to understand plant community assembly in the tropical montane forests of the Andes it is too simple to focus just on environmental sorting by elevational climatic gradients.     

Hubbell's fundamental biodiversity parameter and the Simpson diversity index

Central to Hubbell's neutral theory of biodiversity is a universal, dimensionless fundamental biodiversity parameter that is the product of community size and speciation rate. One of the most important discoveries of Hubbell's theory is that the species‐abundance distribution and the species–area relationship of the neutral metacommunity is completely determined by this fundamental biodiversity parameter, although the diversity patterns of the local community are collectively determined by the biodiversity parameter and migration. Using the relative abundance of species and following the concept of heterozygosity of population genetics, here we developed an analytical relationship between this biodiversity parameter and the well‐known Simpson diversity index. This relationship helps bridge the evolutionary aspect of biodiversity to the ecological and statistical aspect of the diversity. The relationship between these two parameters suggests that diversity patterns of the metacommunity can also be equally described by the Simpson index. This relationship provides an alternative approach to interpret and estimate the fundamental biodiversity parameter for the metacommunity.     

Measuring and modeling dispersal of adult zooplankton

Habitat fragmentation poses an inherent problem for metacommunity dynamics, as dispersal among communities is hindered by increasing isolation and the loss of patches. Wetlands are one such system that have undergone excessive destruction and fragmentation in recent years. Zooplankton within these communities have historically been considered frequent and widespread dispersers, but direct, quantitative measures of zooplankton dispersal are rare. In this study, I performed two experiments to quantify zooplankton dispersal and to identify the primary dispersal vectors. I first set up an array of traps at 10, 30, 60, 120 and 180 m around an isolated pond to collect dispersing individuals. Nearly 1,500 adult zooplankton were captured in traps up to 180 m from the pond, with approximately 60% of dispersers being captured in traps at 10 m from the pond. A second experiment using open and animal-excluded traps suggested that large animals were the primary dispersal vector for these zooplankton. Using a subset of these data, I fit four models to describe the shape and magnitude of adult cladoceran dispersal at this site. All models showed the majority of cladocerans were deposited very close to the source pond, with three models suggesting that the trapping area encompassed 67% or more of the dispersal distances. These results suggest that adult zooplankton movement among ponds may be significant in areas where aquatic habitats are plentiful. Yet, in recent years climate change and anthropogenic disturbances have reduced the number and size of aquatic habitats in many regions of the world, likely curtailing effective transport of individuals in many cases. As a result, fragmented zooplankton metacommunities may experience increased dispersal limitation, stronger priority effects, higher levels of inbreeding and selection against traits engendering high dispersability. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

How does adaptive consumer movement affect population dynamics in consumer-resource metacommunities with homogeneous patches?

This article uses simple models to explore the impact of adaptive movement by consumers on the population dynamics of a consumer-resource metacommunity consisting of two identical patches. Consumer-resource interactions within a patch are described by the Rosenzweig-MacArthur predator-prey model, and these dynamics are assumed to be cyclic in the absence of movement. The per capita movement rate from one patch to the other is an increasing function of the difference between the per capita birth minus death rate in the destination patch and that in the currently occupied patch. Several variations on this model are considered. Results show that adaptive movement frequently creates anti-phase cycles in the two patches; these suppress the predator-prey cycle and lead to low temporal variation of the total population sizes of both species. Paradoxically, even when movement is very sensitive to the fitness difference between patches, perfect synchrony of patches is often much less likely than in comparable systems with random movement. Under these circumstances adaptive movement of consumers often generates differences in the average properties of the two patches. In addition, mean global densities and responses to global perturbations often differ greatly from similar systems with no movement or random movement.     

Dispersal-mediated coexistence of competing predators

Models of metapopulations have often ignored local community dynamics and spatial heterogeneity among patches. However, persistence of a community as a whole depends both on the local interactions and the rates of dispersal between patches. We study a mathematical model of a metacommunity with two consumers exploiting a resource in a habitat of two different patches. They are the exploitative competitors or the competing predators indirectly competing through depletion of the shared resource. We show that they can potentially coexist, even if one species is sufficiently inferior to be driven extinct in both patches in isolation, when these patches are connected through diffusive dispersal. Thus, dispersal can mediate coexistence of competitors, even if both patches are local sinks for one species because of the interactions with the other species. The spatial asynchrony and the competition-colonization trade-off are usual mechanisms to facilitate regional coexistence. However, in our case, two consumers can coexist either in synchronous oscillation between patches or in equilibrium. The higher dispersal rate of the superior prompts rather than suppresses the inferior. Since differences in the carrying capacity between two patches generate flows from the more productive patch to the less productive, loss of the superior by emigration relaxes competition in the former, and depletion of the resource by subsidized consumers decouples the local community in the latter.     

Neutral and non-neutral macroecology

Because of the multiscalar nature of processes underlying biodiversity dynamics, macroecology has emerged as a discipline that seeks to build an understanding of this complexity by examining statistical patterns in large assemblages of species in geographic space and ecological time. Models that assume individual organisms within trophically defined assemblages are ecologically equivalent can produce many patterns identified by macroecology. Neutral models predict two important dynamical patterns that can be tested in real assemblages. First, they predict that species diversity will decline within an assemblage over time. The rate of this decay in species diversity can be predicted from estimates of migration rates from a “metacommunity” or species pool. Second, neutral models predict a divergence of species composition among local communities over time. The rate and degree of divergence among communities also depend on the migration rate. The few studies that have been done to date imply that the rate of migration in real species assemblages is much lower than that required to explain the degree of community similarity maintained in space and time. There are at least two alternative ways to extend neutral models to incorporate more biological realism. First, competitive asymmetries among species may be introduced to allow for the possibility that individuals of some species may have an advantage in replacing individuals that die. Second, environmental heterogeneity can be introduced by assuming sites available to individuals differ in quality to individuals of different species. The neutral model, because of its conceptual simplicity and rigor, should be considered as a null model for baseline comparison to actual patterns of distribution, abundance, species composition, and beta diversity.

Zusammenfassung

Wegen der multiskalaren Natur der Prozesse, die der Biodiversitätsdynamik zugrunde liegen, entstand die Makroökologie als eine Disziplin, die anstrebt ein Verständnis dieser Komplexität zu schaffen, indem sie statistische Muster in großen Vergesellschaftungen von Arten im geografischen Raum und ökologischer Zeit untersucht. Modelle, die davon ausgehen, dass individuelle Organismen innerhalb trophisch definierter Vergesellschaftungen ökologisch äquivalent sind, können viele Muster erzeugen, die durch die Makroökologie indentifiziert werden. Neutrale Modelle sagen zwei wichtige dynamische Muster vorher, die in realen Vergesellschaftungen getestet werden können. Als Erstes sagen sie vorher, dass die Artendiversität in einer Vergesellschaftung mit der Zeit abnehmen wird. Die Rate der Abnahme der Artendiversität kann über Schätzungen der Migrationsraten aus einer Metagemeinschaft bzw. einem Artenpool vorhergesagt werden. Als Zweites sagen neutrale Modelle eine Divergenz der Artenzusammensetzung zwischen den lokalen Gemeinschaften mit der Zeit vorher. Die Rate und der Grad der Divergenz zwischen den Gemeinschaften hängt ebenfalls von der Migrationsrate ab. Die wenigen Untersuchungen, die bis heute gemacht wurden, implizieren, dass die Rate der Migration in realen Artenvergesellschaftungen viel geringer als erforderlich sind, um den Grad der Gemeinschaftsähnlichkeit zu erklären, der in Raum und Zeit aufrecht erhalten wird. Es gibt mindestens zwei alternative Weisen neutrale Modelle zu erweitern, um mehr biologische Realität mit einzubeziehen. Als Erstes können Asymmetrien der Konkurrenz unter Arten einbezogen werden, um die Möglichkeit zu zulassen, dass Individuen einiger Arten einen Vorteil bei der Ersetzung von sterbenden Individuen haben. Als Zweites kann die Umweltheterogenität mit einbezogen werden, indem angenommen wird, dass sich die verfügbaren Standorte in ihrer Qualität für Individuen verschiedener Arten unterscheiden. Wegen seiner konzeptuellen Einfachheit und Starrheit sollte das neutrale Modell als Null-Modell für grundlegende Vergleiche von Verbreitung, Abundanz, Artenzusammensetzung und Betadiversität angesehen werden.