Temporal Beta Diversity Research Articles

Bottom–up and top–down effects on plant communities: nutrients limit productivity, but insects determine diversity and composition

Top–down effects of herbivores and bottom–up effects of nutrients shape productivity and diversity across ecosystems, yet their single and combined effects on spatial and temporal beta diversity is unknown. We established a field experiment in which the abundance of insect herbivores (top–down) and soil nitrogen (bottom–up) were manipulated over six years in an existing old‐field community. We tracked plant α and β diversity – within plot richness and among plot biodiversity‐ and aboveground net primary productivity (ANPP) over the course of the experiment. We found that bottom–up factors affected ANPP while top–down factors influenced plant community structure. Across years, while N reduction lowered ANPP by 10%, N reduction did not alter ANPP relative to control plots. Further, N reduction lowered ANPP by 20% relative to N addition plots. On the other hand, the reduction of insect herbivores did not alter plant richness (α diversity) yet consistently promoted Shannon's evenness, relative to plots where insect herbivores were present. Further, insect herbivores promoted spatial‐temporal β diversity. Overall, we found that the relative importance of top–down and bottom–up controls of plant ANPP, plant α diversity, and composition (β diversity) can vary significantly in magnitude and direction. In addition, their effects varied through time, with bottom–up effects influencing ANPP quickly while the effects of top–down factors emerging only late in the experiment to influence plant community composition via shifts in plant dominance.

Thirty-year recovery of mollusc communities after nuclear experimentations on Fangataufa atoll (Tuamotu, French Polynesia).

A 30-year study of temporal changes in gastropod community structure on the reefs of a Pacific Ocean atoll (Fangataufa, Tuamotu Archipelago) subjected to atmospheric nuclear tests during the 1960s offered the opportunity for an otherwise impossible field experiment that could help ecologists understand mollusc primary succession. Reef molluscs were partly or entirely wiped out by the heat of the nuclear tests and the reefs were recolonized by ocean larvae. On all reefs, community composition before the tests was very different from what it evolved to afterwards. A new method of analysis was developed to study the temporal variation in community composition before versus after the tests (temporal beta diversity). Analyses showed that community compositions diverged through time among the reefs. Only some species can survive the harsh conditions of supralittoral zones, so the same species recolonized them; environmental filtering controlled the development of the new communities. In the reef flat and edge zones, differences in community composition seem to be the result of neutral stochastic colonization by larvae coming from the open ocean. All reefs developed a community composition quite different from that before the nuclear tests.

Temporal Beta Diversity of Bird Assemblages in Agricultural Landscapes: Land Cover Change vs. Stochastic Processes.

Temporal variation in the composition of species assemblages could be the result of deterministic processes driven by environmental change and/or stochastic processes of colonization and local extinction. Here, we analyzed the relative roles of deterministic and stochastic processes on bird assemblages in an agricultural landscape of southwestern France. We first assessed the impact of land cover change that occurred between 1982 and 2007 on (i) the species composition (presence/absence) of bird assemblages and (ii) the spatial pattern of taxonomic beta diversity. We also compared the observed temporal change of bird assemblages with a null model accounting for the effect of stochastic dynamics on temporal beta diversity. Temporal assemblage dissimilarity was partitioned into two separate components, accounting for the replacement of species (i.e. turnover) and for the nested species losses (or gains) from one time to the other (i.e. nestedness-resultant dissimilarity), respectively. Neither the turnover nor the nestedness-resultant components of temporal variation were accurately explained by any of the measured variables accounting for land cover change (r2<0.06 in all cases). Additionally, the amount of spatial assemblage heterogeneity in the region did not significantly change between 1982 and 2007, and site-specific observed temporal dissimilarities were larger than null expectations in only 1% of sites for temporal turnover and 13% of sites for nestedness-resultant dissimilarity. Taken together, our results suggest that land cover change in this agricultural landscape had little impact on temporal beta diversity of bird assemblages. Although other unmeasured deterministic process could be driving the observed patterns, it is also possible that the observed changes in presence/absence species composition of local bird assemblages might be the consequence of stochastic processes in which species populations appeared and disappeared from specific localities in a random-like way. Our results might be case-specific, but if stochastic dynamics are generally dominant, the ability of correlative and mechanistic models to predict land cover change effects on species composition would be compromised.

A novel multiple-site extension to pairwise partitioned taxonomic beta diversity

Our understanding of the causes of variation in taxonomic composition, or beta diversity, is progressing rapidly, thanks in part to recent methodological advances. For example, methods for partitioning beta diversity into its “replacement” and “richness” components have helped reveal patterns that had been undetected by traditional analyses. These partitioning methods are derived from pairwise dissimilarity measures, and are thus well suited to many conventional beta diversity analyses, including “distance decay” relationships. However, pairwise beta diversity measures have limitations, including their lack of information about taxa that are shared among three or more sites. Recently, a new suite of multiple-site counterparts to the pairwise partitioning measures of beta diversity was proposed, but the pairwise analogs upon which these were based were subsequently criticized, and compelling arguments were presented in favor of other partitioning approaches. Here, we introduce multiple-site partitioning measures that address these shortcomings, and illustrate their desirable properties using numerical simulations. We also provide an empirical example of their utility by analyzing the temporal beta diversity of breeding birds within the conterminous USA. We show that temporal beta diversity is predominantly driven by replacement rather than richness differences, and correspondingly, that correlations between temporal beta diversity and productivity and elevation are driven primarily by the replacement component. Furthermore, in contrast to existing multiple-site measures, we show that richness differences do play an important part in driving overall beta diversity patterns. Our new multiple-site measures therefore complement existing methods for analyzing beta diversity, and are especially suitable when compositional heterogeneity is the response of interest.

Effect of (a)synchronous light fluctuation on diversity, functional and structural stability of a marine phytoplankton metacommunity

Disentangling the mechanisms that maintain the stability of communities and ecosystem properties has become a major research focus in ecology in the face of anthropogenic environmental change. Dispersal plays a pivotal role in maintaining diversity in spatially subdivided communities, but only a few experiments have simultaneously investigated how dispersal and environmental fluctuation affect community dynamics and ecosystem stability. We performed an experimental study using marine phytoplankton species as model organisms to test these mechanisms in a metacommunity context. We established three levels of dispersal and exposed the phytoplankton to fluctuating light levels, where fluctuations were either spatially asynchronous or synchronous across patches of the metacommunity. Dispersal had no effect on diversity and ecosystem function (biomass), while light fluctuations affected both evenness and community biomass. The temporal variability of community biomass was reduced by fluctuating light and temporal beta diversity was influenced interactively by dispersal and fluctuation, whereas spatial variability in community biomass and beta diversity were barely affected by treatments. Along the establishing gradient of species richness and dominance, community biomass increased but temporal variability of biomass decreased, thus highest stability was associated with species-rich but highly uneven communities and less influenced by compensatory dynamics. In conclusion, both specific traits (dominance) and diversity (richness) affected the stability of metacommunities under fluctuating conditions.

Statistical methods for temporal and space–time analysis of community composition data

This review focuses on the analysis of temporal beta diversity, which is the variation in community composition along time in a study area. Temporal beta diversity is measured by the variance of the multivariate community composition time series and that variance can be partitioned using appropriate statistical methods. Some of these methods are classical, such as simple or canonical ordination, whereas others are recent, including the methods of temporal eigenfunction analysis developed for multiscale exploration (i.e. addressing several scales of variation) of univariate or multivariate response data, reviewed, to our knowledge for the first time in this review. These methods are illustrated with ecological data from 13 years of benthic surveys in Chesapeake Bay, USA. The following methods are applied to the Chesapeake data: distance-based Moran's eigenvector maps, asymmetric eigenvector maps, scalogram, variation partitioning, multivariate correlogram, multivariate regression tree, and two-way MANOVA to study temporal and space-time variability. Local (temporal) contributions to beta diversity (LCBD indices) are computed and analysed graphically and by regression against environmental variables, and the role of species in determining the LCBD values is analysed by correlation analysis. A tutorial detailing the analyses in the R language is provided in an appendix.

Ground spider assemblages (Araneae: Gnaphosidae) along an urban-rural gradient in the city of Heraklion, Greece

The responses of gnaphosid spiders to an urbanization gradient (urban-suburban-rural areas) were studied using pitfall traps in and near the city of Heraklion, in Crete, Greece, from October 2010 to October 2011. Our results indicate that richness and abundance of species of spiders decreased from the rural area to the center of the city, but not significantly so. Temporal beta diversity was significantly higher in urban areas, which indicates a high temporal variability in species composition of gnaphosid assemblages in the center of the city that takes the form of complementary rather than synchronized phenologies of co-occurring species, as expected in habitats degraded by high levels of disturbance. There was no specific pattern in the body size distribution of the gnaphosids along the urban-rural gradient, and thus the decrease in mean body size with increase in urbanization hypothesis was not supported by our results. However, the species composition recorded in urban areas was very different from that in suburban and rural areas. The percentage of individuals in the catches that were generalist species differed significantly along the gradient. As generalist gnaphosid species made up 72.01% of the total caught in the center of the city, and only 9.53% and 2.07% in suburban and rural areas respectively, our results support the opportunistic species hypothesis. Our analyses indicate that Urozelotes rusticus, a well known synanthropic species recorded for the first time in Greece, is an indicator of high levels of urbanization in Heraklion.

The response of plant diversity to grazing varies along an elevational gradient

Summary Species richness of plants generally decreases along elevational gradients or peaks at intermediate elevations. Land use including grazing by wild and domestic herbivores also affects plant communities and diversity, but how grazing affects plant diversity along elevational gradients is less clear. Using a field experiment along an elevational gradient in Norway, we tested whether the impact of grazing on plant diversity interacts with elevation. Vascular plant communities were sampled following 10 years of experimental sheep grazing (decreased/ceased, maintained and increased densities of 0, 25 and 80 sheep km−2) and compared with pre‐experimental baseline data. The impact of grazing on species richness (alpha diversity) was investigated at the species level, and temporal community variance was used to investigate community stability (temporal beta diversity). Spatial community variance (spatial beta diversity) at the enclosure level was used to test whether community homogenization differed between treatments. Species richness and temporal stability varied between treatments and along the elevational gradient. Where grazing was ceased, species richness declined by up 3.7 species at low elevations and increased by up to 3.5 species at high elevations, whilst where grazing was maintained or increased, changes were less extreme along the elevational gradient. Temporal stability of plant communities was highest at low elevations and lowest at high elevations where grazing was reduced. There were no clear differences in spatial homogenization between grazing treatments, although spatial species turnover increased in heathlands where grazing decreased. Synthesis. This study shows that the effect of grazing on plant diversity varies with elevation, and grazing herbivores can thus directly affect elevational patterns of plant diversity. Grazing can buffer changes in plant communities and species richness, even in the face of other environmental drivers such as climatic warming.

Ground beetles respond to grazing at assemblage level, rather than species-specifically: the case of Cretan shrublands

Phrygana, maquis and subalpine shrublands are the most typical ecosystem types on the island of Crete. However, these formations are mostly mismanaged and poorly studied in terms of their fauna and the effect of long term land-use. We studied the impact of goat and sheep grazing on the diversity patterns of carabid beetles at 40 sites in relation to possible habitat uniqueness and geographic zonation. In total, 19,322 individuals belonging to 42 Carabidae species were identified. The composition of site assemblages was mainly determined by altitude. The six Cretan endemic species collected were widely distributed on the island and most of them were dominant. Species richness and equitability were significantly lower at overgrazed sites compared to areas with low grazing intensity. The diversity patterns of carabids were influenced by the level of grazing in each study area. As species richness and diversity were maximal under moderate/relatively high levels of grazing (hump-shaped pattern), our results support the intermediate disturbance hypothesis. Low tolerance of carabid taxa to intensive grazing was also ascertained through the significantly high values of temporal beta diversity at overgrazed sites, independent of local species richness. We conclude that Carabidae are good indicators of grazing pressure in Crete at assemblage level, rather than species-specifically. To maintain diversity and enhance niche space of invertebrate fauna, the spatiotemporal discontinuity of grazing is essential.

Rates of Species Accumulation and Taxonomic Diversification during Phototrophic Biofilm Development Are Controlled by both Nutrient Supply and Current Velocity

The accumulation of new and taxonomically diverse species is a marked feature of community development, but the role of the environment in this process is not well understood. To address this problem, we subjected periphyton in laboratory streams to low (10-cm · s(-1)), high (30-cm · s(-1)), and variable (9- to 32-cm · s(-1)) current velocity and low- versus high-nutrient inputs. We examined how current velocity and resource supply constrained (i) the rates of species accumulation, a measure of temporal beta-diversity, and (ii) the rates of diversification of higher taxonomic categories, defined here as the rate of higher taxon richness increase with the increase of species richness. Temporal biofilm dynamics were controlled by a strong nutrient-current interaction. Nutrients accelerated the rates of accumulation of new species, when flow velocity was not too stressful. Species were more taxonomically diverse under variable than under low-flow conditions, indicating that flow heterogeneity increased the niche diversity in the high-nutrient treatments. Conversely, the lower diversification rates under high- than under low-nutrient conditions at low velocity are explained with finer resource partitioning among species, belonging to a limited number of related genera. The overall low rates of diversification in high-current treatments suggest that the ability to withstand current stress was conserved within closely related species. Temporal heterogeneity of disturbance has been shown to promote species richness, but here we further demonstrate that it also affects two other components of biodiversity, i.e., temporal beta-diversity and diversification rate. Therefore, management efforts for preserving the inherent temporal heterogeneity of natural ecosystems will have detectable positive effects on biodiversity.

Projected impacts of climate change on spatio‐temporal patterns of freshwater fish beta diversity: a deconstructing approach

ABSTRACTAim To assess the potential impacts of future climate change on spatio‐temporal patterns of freshwater fish beta diversity.Location Adour–Garonne River Basin (France).Methods We first applied an ensemble modelling approach to project annually the future distribution of 18 fish species for the 2010–2100 period on 50 sites. We then explored the spatial and temporal patterns of beta diversity by distinguishing between its two additive components, namely species turnover and nestedness.Results Taxonomic homogenization of fish assemblages was projected to increase linearly over the 21st century, especially in the downstream parts of the river gradient. This homogenization process was almost entirely caused by a decrease in spatial species turnover. When considering the temporal dimension of beta diversity, our results reveal an overall pattern of decreasing beta diversity along the upstream–downstream river gradient. In contrast, when considering the turnover and nestedness components of temporal beta diversity we found significant U‐shaped and hump‐shaped relationships, respectively.Main conclusions Future climate change is projected to modify the taxonomic composition of freshwater fish assemblages by increasing their overall similarity over the Adour–Garonne River Basin. Our findings suggest that the distinction between the nestedness and turnover components of beta diversity is not only crucial for understanding the processes shaping spatial beta‐diversity patterns but also for identifying localities where the rates of species replacement are projected to be greatest. Specifically we recommend that future conservation studies should not only consider the spatial component of beta diversity but also its dynamic caused by climate warming.

Instability of Copronecrophagous Beetle Assemblages (Coleoptera: Scarabaeinae) in a Mountainous Tropical Landscape of Mexico

We analyzed changes over time in species composition and functional guild structure (temporal beta diversity) for natural assemblages and those modified by humans in a fragmented, tropical mountain landscape. The assemblages belong to cloud forests (the original vegetation type), secondary forests, traditional shaded coffee plantations, commercial shaded coffee plantations, and a cattle pasture. Copronecrophagous beetles, subfamily Scarabaeinae (Insecta: Coleoptera: Scarabaeidae), were used as the indicator group. This group has been used in previous studies and other tropical forests and has been found to be a good indicator of the effects of anthropogenic change. For each assemblage, we compared samples that were collected several years apart. Changes were found in species composition, order of abundance, and in the proportion that a given species is present in the different functional groups. The changes that occurred between samplings affected the less abundant species in the cloud forest and in the pasture. In the other vegetation types, both abundant and less abundant species were affected. Their order of abundance and proportion in the different guilds also changed. This study shows that, although landscape richness remains relatively constant, richness at the local level (alpha diversity) changes notably even over short lapses of time. This could be a characteristic of landscapes with intermediate degrees of disturbance (such as those that have been partially modified for human use), where assemblage composition is very fluid.

Species richness in Mediterranean agroecosystems: Spatial and temporal analysis for biodiversity conservation

We used dung beetles as an indicator group to identify the most important habitats for biodiversity conservation in a Mediterranean traditional agroecosystem. The relationships between traditional grazing and farming activities and biodiversity were analysed by comparing species richness and temporal turnover across three different habitat types, defined according to vegetation structure and human land use. In this study, both spatial (landscape, among-habitats) and temporal (intra-annual, seasonal) analyses were contemplated at mesoscale. The measurement and use of temporal beta diversity (i.e. an index of temporal species turnover) in conservation biology showed that it is a simple method to characterize and compare different habitat species-assemblages, particularly when time seems to be a significant factor in explaining biodiversity features. Our results showed open mosaic areas as the richest and most temporally heterogeneous habitats. We suggest the maintenance of traditional human activities carried out in these areas, since they have been a significant diversification agent, to avoid the loss of the high Mediterranean biological diversity.