Hierarchical Dynamics of Ecological Communities: Do Scales of Space and Time Match?

Theory posits that community dynamics organize at distinct hierarchical scales of space and time, and that the spatial and temporal patterns at each scale are commensurate. Here we use time series modeling to investigate fluctuation frequencies of species groups within invertebrate metacommunities in 26 boreal lakes over a 20-year period, and variance partitioning analysis to study whether species groups with different fluctuation patterns show spatial signals that are commensurate with the scale-specific fluctuation patterns identified. We identified two groups of invertebrates representing hierarchically organized temporal dynamics: one species group showed temporal variability at decadal scales (slow patterns of change), whilst another group showed fluctuations at 3 to 5-year intervals (faster change). This pattern was consistently found across all lakes studied. A spatial signal was evident in the slow but not faster-changing species groups. As expected, the spatial signal for the slow-changing group coincided with broad-scale spatial patterns that could be explained with historical biogeography (ecoregion delineation, and dispersal limitation assessed through a dispersal trait analysis). In addition to spatial factors, the slow-changing groups correlated with environmental variables, supporting the conjecture that boreal lakes are undergoing environmental change. Taken together our results suggest that regionally distinct sets of taxa, separated by biogeographical boundaries, responded similarly to broad-scale environmental change. Not only does our approach allow testing theory about hierarchically structured space-time patterns; more generally, it allows assessing the relative role of the ability of communities to track environmental change and dispersal constraints limiting community structure and biodiversity at macroecological scales.     

Influence of nutrients, submerged macrophytes and zooplankton grazing on phytoplankton biomass and diversity along a latitudinal gradient in Europe

In order to evaluate latitudinal differences in the relationship of phytoplankton biomass and diversity with environmental conditions in shallow lakes, we sampled 98 shallow lakes from three European regions: Denmark (DK), Belgium/The Netherlands (BNL) and southern Spain (SP). Phytoplankton biomass increased with total phosphorus (TP) concentrations and decreased with submerged macrophyte cover across the three regions. Generic richness was significantly negatively related to submerged macrophyte cover and related environmental variables. Zooplankton:phytoplankton biomass ratios were positively related to submerged macrophyte cover and negatively to phytoplankton generic richness in DK and BNL, suggesting that the low generic richness in lakes with submerged macrophytes was due to a higher zooplankton grazing pressure in these regions. In SP, phytoplankton generic richness was not influenced by zooplankton grazing pressure but related to conductivity. We observed no relationship between phytoplankton generic richness and TP concentration in any of the three regions. The three regions differed significantly with respect to mean local and regional generic richness, with BNL being more diverse than the other two regions. Our observations suggest that phytoplankton diversity in European shallow lakes is influenced by submerged macrophyte cover indirectly by modulating zooplankton grazing. This influence of submerged macrophytes and zooplankton grazing on phytoplankton diversity decreases from north to south.     

Patterns of temporal community turnover are spatially synchronous across boreal lakes

1. Ecosystems are often exposed to broad‐scale environmental change, which can potentially synchronise community dynamics and biodiversity trends. Detection of temporal coherence may, however, depend on the metrics used and their sensitivity to detect change, requiring several lines of evidence to elucidate the full range of temporal responses to environmental change. 2. Here, we tested whether the patterns of synchrony among littoral invertebrate communities of Swedish lakes over 20 years (1988–2007) differed when analysed using univariate (taxon richness, evenness, Shannon diversity and total abundance) or multivariate (temporal turnover in community composition) metrics. We included both culturally acidified and circumneutral lakes to examine whether anthropogenic stress influenced the patterns of synchrony. 3. Average total abundance, taxon richness and temporal turnover in community composition changed monotonically with time, while evenness and Shannon diversity fluctuated around a long‐term mean. However, among‐lake variability was high, resulting in a weak temporal coherence. Only trends of temporal turnover changed synchronously across lakes, irrespective of their acidification history. 4. Spatially synchronous trends in turnover across lakes were correlated with increasing water colour and decreasing sulphate concentrations, showing the importance of regional drivers of spatiotemporal coherence. 5. Our results underpin an increasing body of evidence that the detection of diversity patterns varies among metrics that ignore (taxon richness, evenness, Shannon diversity) or consider (turnover) species identities. More generally, our results suggest that community‐level studies of synchrony are suitable for elucidating the role of intrinsic versus extrinsic factors in mediating complex community assembly processes in the long term. This, in turn, contributes to our understanding of temporal patterns of biodiversity.     

Fish diversity in European lakes: geographical factors dominate over anthropogenic pressures

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Phytoplankton diversity (alpha,beta, and gamma) from the Araguaia River tropical floodplain lakes (central Brazil)

Alpha, beta and gamma are three components of species diversity. Knowing these attributes in floodplain lake phytoplankton communities is vital when selecting conservation areas. Species diversity is commonly used with other taxonomic groups, but rarely with phytoplankton. We compared the number of phytoplankton species (alpha diversity) from 21 Middle Araguaia River floodplain lakes in the 2000 and 2001 rainy and dry seasons. From these samples we estimated complete survey species richness (gamma diversity), quantified differences in species composition between lakes (beta diversity) and assessed the influence of abiotic variables on beta diversity. We recorded a total of 577 taxa. The Sjack1 estimator indicated that 62.31% of taxa were sampled in the 2000 rainy and 67.65% dry seasons, and 68.36% in the 2001 rainy and 73.5% dry seasons. In almost all seasons, alpha diversity negatively correlated with latitude. Beta diversity (β-1) was higher in high water periods, especially in 2000. This may have been caused by isolated heavy rainfall, which would have increased environmental heterogeneity and raised beta diversity. DCA showed differences in phytoplankton composition between rainy and dry seasons in 2000 and 2001, reflecting the influence of flood pulses on phytoplankton composition. The Mantel test indicated spatial distribution patterns where geographically more distant lakes had less-similar phytoplankton communities. Handling editor: J. Padisak     

Environmental and historical effects on richness and endemism patterns of carabid beetles in the western Palaearctic

Identifying determinants of spatial diversity gradients is in the focus of biodiversity-related research and has gained considerable importance regarding global change and conservation strategies. Despite their overwhelming diversity and the crucial role of invertebrates in ecosystem functioning, our understanding of factors driving broad-scale invertebrate diversity is limited. Tackling this issue, our study analyzes macroecological patterns of a highly diverse insect taxon across large parts of the western Palaearctic. We used regression modelling to assess the influence of environmental factors on overall, widespread and restricted-range (endemic) carabid beetle diversity. Single-term regressions and variation partitioning among climatic, topographic and spatial variables showed that total carabid diversity as well as richness patterns of widespread species were most strongly correlated with spatially structured variables related to current climate (measures of ambient energy and, to a lesser degree, precipitation and AET). In contrast, restricted-range (endemic) species were most notably related to range in elevation. We discuss the possible role of this factor as a surrogate measure of historical processes and the impact of history on contemporary diversity distributions. Our results indicate that while overall diversity patterns of carabids strongly reflect current climate conditions, this primarily reflects the more widespread species, whereas the spatial distribution of restricted-range species is still significantly affected by historical processes. Thus, for a general understanding of determinants and mechanisms of broad-scale diversity, taking into account dispersal abilities and range sizes of species is essential, especially as large parts of global biodiversity are represented by invertebrates with low dispersal powers.     

Environmental heterogeneity among lakes promotes hyper β‐diversity across phytoplankton communities

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Diversity of Eastern North American Ant Communities along Environmental Gradients

Studies of species diversity patterns across regional environmental gradients seldom consider the impact of habitat type on within-site (alpha) and between-site (beta) diversity. This study is designed to identify the influence of habitat type across geographic and environmental space, on local patterns of species richness and regional turnover patterns of ant diversity in the northeastern United States. Specifically, I aim to 1) compare local species richness in paired open and forested transects and identify the environmental variables that best correlate with richness; and 2) document patterns of beta diversity throughout the region in both open and forested habitat. I systematically sampled ants at 67 sites from May to August 2010, spanning 10 degrees of latitude, and 1000 meters of elevation. Patterns of alpha and beta diversity across the region and along environmental gradients differed between forested and open habitats. Local species richness was higher in the low elevation and warmest sites and was always higher in open habitat than in forest habitat transects. Richness decreased as temperature decreased or elevation increased. Forested transects show strong patterns of decreasing dissimilarity in species composition between sites along the temperature gradient but open habitat transects did not. Maximum temperature of the warmest month better predicted species richness than either latitude or elevation. I find that using environmental variables as key predictors of richness yields more biologically relevant results, and produces simpler macroecological models than commonly used models which use only latitude and elevation as predictors of richness and diversity patterns. This study contributes to the understanding of mechanisms that structure the communities of important terrestrial arthropods which are likely to be influenced by climatic change.     

Concordance among assemblages of upland Amazonian lakes and the structuring role of spatial and environmental factors

Conservation planning and biodiversity monitoring rely exclusively on the use of surrogate groups. However, the effectiveness of these groups in representing unknown biodiversity is rarely tested. These tests can be based on cross-taxon concordance (or congruence) analysis. In a second step, the main mechanisms underlying concordance can be investigated. Here, we evaluated the relative influence of local environmental factors and spatial processes on the patterns of beta-diversity exhibited by phytoplankton, zooplankton and benthic invertebrates and the strength of assemblage concordance among these assemblages in 38 upland Amazonian lakes (northern Brazil) over four years. Zooplankton and benthic invertebrates responded exclusively to environmental variables. These results are in accordance with the expected under the species sorting model, where local factors are the main mechanisms shaping assemblage structure. In general, patterns of concordance between the assemblages were weak and varied through time, indicating that the use of surrogates in freshwater ecosystems may be a flawed approach.     

Beta diversity of aquatic invertebrates increases along an altitudinal gradient in a Neotropical mountain

Mountains harbor rich biodiversity and high levels of endemism, particularly due to changes in environmental conditions over short spatial distances, which affects species distribution and composition. Studies on mountain ecosystems are increasingly needed, as mountains are highly threatened despite providing ecosystem services, such as water supply for half of the human population. We aimed to understand the patterns and drivers of alpha and beta diversities of aquatic invertebrates in headwater streams along an altitudinal gradient in the second largest South American mountain range, the Espinhaço mountains. Headwater streams were selected at each 100 m of elevation along an altitudinal gradient ranging from 800 to 1400 m asl, where three substrate types per stream were sampled: leaf litter, gravel, and cobbles. Environmental variables were sampled to represent local riparian canopy cover, instream physical habitat, water quality, climatic data, and land use. Generalized linear models and mixed models were used to test relationships between altitude and the richness and abundance of invertebrates and to assess the influence of environmental variables on the same metrics. Patterns of spatial variation in aquatic invertebrate assemblages along the altitudinal gradient were assessed using multiplicative beta diversity partitioning. The richness and abundance of aquatic invertebrates decreased with increasing altitude, whereas beta diversity increased with increasing altitude. Significant differences in assemblage composition and in relative abundance of invertebrates were observed for both substrates and altitude. We thus show that the high regional beta diversity in aquatic ecosystems in the studied site is due to the high turnover among areas. Abstract in Portuguese is available with online material.     

Selecting networks of nature reserves: methods do affect the long-term outcome

Data on vascular plants of boreal lakes in Finland were used to compare the efficiency of reserve selection methods in representing four aspects of biodiversity over a 63 year period. These aspects included species richness, phylogenetic diversity, restricted range diversity and threatened species. Our results show that the efficiency of reserve selection methods depends on the selection criteria used and on the aspect of biodiversity under consideration. Heuristic methods and optimizing algorithms were nearly equally efficient in selecting lake networks over a small geographical range. In addition, a scoring procedure was observed to be efficient in maintaining different aspects of biodiversity over time. However, the random selection of lakes seems to be the most inefficient option for a reserve network. In general, reserve selection methods seem to favour lakes that maximize one aspect of diversity at the time of selection, but the network may not be the best option for maintaining the maximum diversity over time. The reserve selection methods do affect the long-term outcome but it is impossible to recommend one method over the others unequivocally.     

Congruence,but no cascade—Pelagic biodiversity across three trophic levels in Nordic lakes

Covariation in species richness and community structure across taxonomical groups (cross‐taxon congruence) has practical consequences for the identification of biodiversity surrogates and proxies, as well as theoretical ramifications for understanding the mechanisms maintaining and sustaining biodiversity. We found there to exist a high cross‐taxon congruence between phytoplankton, zooplankton, and fish in 73 large Scandinavian lakes across a 750 km longitudinal transect. The fraction of the total diversity variation explained by local environment alone was small for all trophic levels while a substantial fraction could be explained by spatial gradient variables. Almost half of the explained variation could not be resolved between local and spatial factors, possibly due to confounding issues between longitude and landscape productivity. There is strong consensus that the longitudinal gradient found in the regional fish community results from postglacial dispersal limitations, while there is much less evidence for the species richness and community structure gradients at lower trophic levels being directly affected by dispersal limitation over the same time scale. We found strong support for bidirectional interactions between fish and zooplankton species richness, while corresponding interactions between phytoplankton and zooplankton richness were much weaker. Both the weakening of the linkage at lower trophic levels and the bidirectional nature of the interaction indicates that the underlying mechanism must be qualitatively different from a trophic cascade.     

Ephemeral ecological speciation and the latitudinal biodiversity gradient

The richness of biodiversity in the tropics compared to high‐latitude parts of the world forms one of the most globally conspicuous patterns in biology, and yet few hypotheses aim to explain this phenomenon in terms of explicit microevolutionary mechanisms of speciation and extinction. We link population genetic processes of selection and adaptation to speciation and extinction by way of their interaction with environmental factors to drive global scale macroecological patterns. High‐latitude regions are both cradle and grave with respect to species diversification. In particular, we point to a conceptual equivalence of “environmental harshness” and “hard selection” as eco‐evolutionary drivers of local adaptation and ecological speciation. By describing how ecological speciation likely occurs more readily at high latitudes, with such nascent species especially prone to extinction by fusion, we derive the ephemeral ecological speciation hypothesis as an integrative mechanistic explanation for latitudinal gradients in species turnover and the net accumulation of biodiversity.     

Environmental drivers of beta-diversity patterns in New-World birds and mammals

Current macroecological research places great emphasis on patterns of species richness (alpha diversity) and the underlying ecological and evolutionary processes involved in their origin and maintenance. However, few studies dealing with continental scales have addressed dissimilarities in species composition among areas (beta diversity). Using data for the occurrence of 3836 bird and 1641 mammal species in 4220 cells covering the New World, we assessed whether broad-scale macroecological patterns in beta diversity are related to dissimilarities in environmental variables and biotic units. We employed spatial regression and tree regression to model beta diversity. Difference in altitude was the best predictor of beta diversity. Accordingly, the highest beta diversity values were found in mountainous areas, particularly in the Andes, Central America and western North America. Explanatory variables related to transitions between biotic units (biome, ecoregion) were relatively unimportant. Areas that differ in altitude from their surroundings harbor different sets of species, and this may reflect either species adaptation to particular environmental conditions by range shifts, or species divergence by vicariance, or both.     

The use of long‐term monitoring data for studies of planktonic diversity: a cautionary tale from two Swiss lakes

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Species richness in mammalian herbivores: patterns in the boreal zone

Latitudinal gradients in species diversity are well established for a number of plant and animal taxa. Both historical and present-day environmental factors have been suggested to be responsible for observed patterns. We tested the hypothesis that current environmental features of the environment (primary productivity and regional landscape structure) may explain the longitudinal variation in species richness of mammalian herbivores in the Holarctic boreal zone. Mammalian herbivore species diversity was strongly correlated with a number of environmental variables measured. We reduced the data set by a principal components analysis (PCA), and found that in the Palearctic, species richness is positively related to warm climate (high temperature sum), the number of hardwood species, and the area of boreal forest. In the Nearctic, species richness increases as the length of the growing season and the number of coniferous tree species increase. Thus indirect measures of primary productivity as well as tree species number may accurately predict species richness in mammalian herbivores. In addition, there seems to be a strong species-area effect at the regional level. The larger and more homogeneous in terms of forest coverage a boreal section is, the more species coexist there.     

Productivity-diversity relationships in lake plankton communities

One of the most intriguing environmental gradients connected with variation in diversity is ecosystem productivity. The role of diversity in ecosystems is pivotal, because species richness can be both a cause and a consequence of primary production. However, the mechanisms behind the varying productivity-diversity relationships (PDR) remain poorly understood. Moreover, large-scale studies on PDR across taxa are urgently needed. Here, we examined the relationships between resource supply and phyto-, bacterio-, and zooplankton richness in 100 small boreal lakes. We studied the PDR locally within the drainage systems and regionally across the systems. Second, we studied the relationships between resource availability, species richness, biomass and resource ratio (N:P) in phytoplankton communities using Structural Equation Modeling (SEM) for testing the multivariate hypothesis of PDR. At the local scale, the PDR showed variable patterns ranging from positive linear and unimodal to negative linear relationships for all planktonic groups. At the regional scale, PDRs were significantly linear and positive for phyto- and zooplankton. Phytoplankton richness and the amount of chlorophyll a showed a positive linear relationship indicating that communities consisting of higher number of species were able to produce higher levels of biomass. According to the SEM, phytoplankton biomass was largely related to resource availability, yet there was a pathway via community richness. Finally, we found that species richness at all trophic levels was correlated with several environmental factors, and was also related to richness at the other trophic levels. This study showed that the PDRs in freshwaters show scale-dependency. We also documented that the PDR complies with the multivariate model showing that plant biomass is not mirroring merely the resource availability, but is also influenced by richness. This highlights the need for conserving diversity in order to maintain ecosystem processes in freshwaters.     

Identifying resilience mechanisms to recurrent ecosystem perturbations

The complex nature of ecological systems limits the unambiguous determination of mechanisms that drive resilience to natural disturbance or anthropogenic stress. Using eight-year time series data from boreal lakes with and without bloom formation of an invasive alga (Gonyostomum semen, Raphidophyceae), we studied resilience of phytoplankton communities in relation to recurring bloom impacts. We first characterized phytoplankton community dynamics in both lake types using univariate metrics of community structure (evenness, species richness, biovolume and Simpson diversity). All metrics, except species richness, were substantially altered and showed an inherent stronger variability in bloom lakes relative to reference lakes. We assessed resilience mechanisms using a multivariate time series modelling technique. The models captured clear successional dynamics of the phytoplankton communities in all lakes, whereby different groups of species were substituted sequentially over the ice-free period. The models also identified that G. semen impacts in bloom lakes were only manifested within a single species group, not across species groups, highlighting the rapid renewal of the phytoplankton communities upon bloom collapse. These results provide empirical support of the cross-scale resilience model. Cross-scale resilience could provide an explanation for the paradox that similar species richnesses are seen in bloom-forming lakes and reference lakes despite the clear difference between the community features of the two different sets of lakes investigated.