Environmental determinants of freshwater mollusc biodiversity and identification of priority areas for conservation in Mediterranean water courses

This study evaluates which environmental factors influence the biodiversity, distribution and conservation of freshwater mollusc communities in a Mediterranean Biosphere Reserve in the south-western Iberian Peninsula. Habitat features and two biodiversity indices (native species richness and diversity) were evaluated at 109 locations. Environmental gradients were assessed using principal component analysis, which orders the habitat variables along two gradients: headwater characteristics and water availability. According to a canonical correspondence analysis, the main environmental factors related to the distribution of species and community structure were, also, climate and headwater habitat features (precipitation, order, channel width, slope and pH), and heterogeneity and trophic features (IHF index and instream macrophytes cover). Relationships between biodiversity indices and environmental variables were best explained by a regression model incorporating, basically, aridity index and precipitation as the variables that accounted for most of the variance. This study demonstrates that distribution of freshwater molluscs along a highly stressed by drought Mediterranean region mostly depends on the local pool of species and their adaptive patterns to water availability.     

Distinctive life traits and distribution along environmental gradients of dominant and subordinate Mediterranean ant species

For most animal and plant species, life traits strongly affect their species-specific role, function or position within ecological communities. Previous studies on ant communities have mostly focused on the role of dominant species and the outcome of interspecific interactions. However, life traits of ant species have seldom been considered within a community framework. This study (1) analyses life traits related to ecological and behavioural characteristics of dominant and subordinate ant species from 13 sites distributed throughout the Iberian Peninsula, (2) determines how similar the ant species are within each of the two levels of the dominance hierarchy, and (3) establishes the distribution patterns of these different groups of species along environmental gradients. Our results showed that the differences between dominants and subordinates fall into two main categories: resource exploitation and thermal tolerance. Dominant species have more populated colonies and defend food resources more fiercely than subordinates, but they display low tolerance to high temperatures. We have identified different assemblages of species included within each of these two levels in the dominance hierarchy. The distribution of these assemblages varied along the environmental gradient, shifting from dominant Dolichoderinae and cryptic species in moist areas, to dominant Myrmicinae and hot climate specialists mainly in open and hot sites. We have been able to identify a set of life traits of the most common Iberian ant species that has enabled us to characterise groups of dominant and subordinate species. Although certain common features within the groups of both dominants and subordinates always emerge, other different features allow for differentiating subgroups within each of these groups. These different traits allow the different subgroups coping with particular conditions across environmental gradients.     

Associations between fish assemblages and environmental factors for Mediterranean-type rivers during summer

The structure of summer fish assemblages was examined along longitudinal gradients in 31 Mediterranean-type rivers of the middle Guadiana basin (south-west Iberian Peninsula), using data from 157 sites including small streams to deep rivers. An ordination analysis, based on 16 variables, was applied to species presence, using principal component and canonical correspondence analysis. The results for the habitat data were compared with those for the biological data using a Mantel analysis, and the agreement was highly significant. Spatial structure was considered by partitioning the total variability among the environmental and geographical variables. The fish assemblages showed longitudinal zoning during the summer, with species distributed over gradients of habitat size (depth), water quality (current and physico-chemical variables), and cover (substratum and vegetation), according to their adult size and life history. The size of the habitat that remained available in summer had the greatest biological effect, being the most important factor explaining fish species distribution and assemblage structure during this stressing season. Strictly spatial variation was low, but there was still a high residual variation. Habitat associations and life-history strategies are discussed for native and exotic species.     

Biogeographical analysis of species of the tribe Cytiseae (Fabaceae) in the Iberian Peninsula and Balearic Islands

Aim To analyse quantitatively the biogeographical distribution pattern of species of the Cytiseae Bercht. & J. Presl (= Genisteae Benth.) tribe in the Iberian Peninsula and the Balearic Islands, and to identify environmental variables related to the distributional patterns. Location Iberian Peninsula and Balearic Islands, using the 61 administrative provinces as operational geographical units. Methods In order to identify chorotypes (groups of species with similar geographical distribution), we performed a upgma classification based on the similarity index of Baroni‐Urbani & Buser. The method of McCoy et al. [Ecology 67 (1986), 749] enabled us to detect the significant groups and to differentiate them from those groupings that could be generated at random. Logistic regression analyses and environmental gradient analyses (DCA and CCA) were performed in order to find the relationships between the environmental variables and the observed distributional patterns. Results Sixteen chorotypes were obtained between the Cytiseae species of the Iberian Peninsula and the Balearic Islands. The thermal variables showed the greatest influence in species distribution. Specifically, temperatures (maximum, minimum and mean) of the coldest months were associated with the majority of the chorotypes. Main conclusions The species of the Cytiseae tribe were not randomly distributed in space, and can be classified in groups of species with common distributional patterns. The importance of cold tolerance in the distribution of these species, as well as their preference for acidic soils, was demonstrated. Certain general tendencies seem to exist with respect to the distribution of the biota in the Iberian Peninsula, and these seem to be independent of the taxonomic groups involved and to be determined by convergent macroclimatic factors.     

Mollusc communities along upstream–downstream gradients in small coastal basins of the south-western Iberian Peninsula

Freshwater mollusc communities readily respond to various human-induced stressors, and thus are appropriate models for studying the effects of such stressors on the structure and dynamics of stream macroinvertebrate communities. This paper examined the distribution of freshwater molluscs in 35 stream reaches of 18 small coastal basins in the south-western Iberian Peninsula. Using several multivariate techniques, I showed that mollusc distribution mainly responded to gradients in drainage area, water availability, pollution and salinity. Upstream and downstream communities were clearly differentiated, with the former dominated by freshwater species (Bithynia tentaculata, Galba truncatula, Radix balthica, Ancylus fluviatilis and Planorbarius metidjensis) and the latter by species typical of brackish waters (Peringia ulvae and Myosotella myosotis). There was a clear decrease in species richness from upper to lower reaches. The conservation of mollusc communities of these small basins requires a deeper understanding of their ecological requirements, effective control of urban discharges and an analysis of their interaction with invasive species.     

Mollusc and plant assemblages controlled by different ecological gradients at Eastern European fens

Ecological patterns of mollusc assemblages and vegetation in relation to water chemistry, water regime, nutrient availability and climate were studied in eastern Polish lowland fens. Our goal was to examine if major compositional changes differ for molluscs and vegetation under the joint influence of multiple ecological gradients. Altogether 32 fen sites were investigated in 2010–2011, and analyzed using metric multidimensional scaling, cluster analysis and generalized additive models. Two major gradients driving the differences in mollusc species composition were revealed. The main direction of compositional changes was associated with the water table gradient, governing a species turnover from inundated and strongly water-logged sites occupied mostly by aquatic mollusc species, to moderately wet sites with the predominance of fen and meadow species. The second most important gradient for molluscs was that of mineral richness. For vegetation, three major gradients explained the changes in species composition. The highest importance was assigned to the nitrogen-to-phosphorus availability gradient (defined as a shift from N-limited to P-limited vegetation), followed by the water table gradient, and the mineral richness gradient. Our results demonstrate that the impact of mineral richness gradient, which has been often reported as the major determinant of compositional changes of fen molluscs and vegetation, can be exceeded by other ecological gradients of comparable variation. We also document for the first time that the main species turnover of fen vegetation is not accompanied by the analogous change in species composition of mollusc assemblages, due to a different sensitivity of these taxa to particular environmental factors (i.e. water level dynamics and type of nutrient limitation).     

Zooplankton community structure along a pollution gradient at fine geographical scales in river ecosystems: The importance of species sorting over dispersal

The release of anthropogenic pollution into freshwater ecosystems has largely transformed biodiversity and its geographical distribution patterns globally. However, for many communities including ecologically crucial ones such as zooplankton, it is largely unknown how different communities respond to environmental pollution. Collectively, dispersal and species sorting are two competing processes in determining the structure and geographical distribution of zooplankton communities in running water ecosystems such as rivers. At fine geographical scales, dispersal is usually considered as the dominant factor; however, the relative role of species sorting has not been evaluated well, mainly because significant environmental gradients rarely exist along continuously flowing rivers. The Chaobai River in northern China represents a rare system, where a significant environmental gradient exists at fine scales. Here, we employed high‐throughput sequencing to characterize complex zooplankton communities collected from the Chaobai River, and tested the relative roles of dispersal and species sorting in determining zooplankton community structure along the pollution gradient. Our results showed distinct patterns of zooplankton communities along the environmental gradient, and chemical pollutant‐related factors such as total phosphorus and chlorophyll‐a were identified as the major drivers for the observed patterns. Further partial redundancy analyses showed that species sorting overrode the effect of dispersal to shape local zooplankton community structure. Thus, our results reject the dispersal hypothesis and support the concept that species sorting caused by local pollution can largely determine the zooplankton community structure when significant environmental gradients exist at fine geographical scales in highly polluted running water ecosystems.     

Complex patterns of environmental niche evolution in Iberian columbines (genus Aquilegia,Ranunculaceae)

Aims This study explores the patterns of niche differentiation in a group of seven closely related columbines (genus Aquilegia, Ranunculaceae) from the Iberian Peninsula. Populations of these columbines are subject to complex patterns of divergent selection across environments, which partly explain the taxonomic structure of the group. This suggests the hypothesis that niche divergence must have occurred along the process of diversification of the group.Methods We used MaxEnt to build environmental niche models of seven subspecies belonging to the three species of Aquilegia present in the Iberian Peninsula. From these models, we compared the environmental niches through two different approaches: ENMtools and multivariate methods.Important findings MaxEnt distributions conformed closely to the actual distribution of the study taxa. ENMtools methods failed to uncover any clear patterns of niche differentiation or conservatism in Iberian columbines. Multivariate analyses indicate the existence of differentiation along altitudinal gradients and along a gradient of climatic conditions determined by the summer precipitation and temperatures. However, climatic conditions related to winter temperature and precipitation, as well as soil properties, were equally likely to show conservatism or divergence. The complex patterns of niche evolution we found suggest that Iberian Columbines have not been significantly constrained by forces of niche conservatism, so they could respond adaptively to the fast and profound climate changes in the Iberian Peninsula through the glacial cycles of the Pleistocene.     

Vegetation composition and its relationship with the environment in mallines of north Patagonia,Argentina

“Mallines” are characteristic Patagonian wet meadows. The objectives of this study were to describe plant community composition in the main mallines in northern Patagonia and to determine the influence of selected environmental variables on the distribution of vegetation. Fifty-two sites were selected for vegetation surveys and measurements of water table (WT) depth, soil pH, electric conductivity (EC), and mean annual precipitation. We performed cluster analysis for vegetation classification and correspondence analysis (CA) and canonical correspondence analysis (CCA) for vegetation ordination. Plant composition was mostly related to both environmental variables and longitude and that it was not possible to disentangle the two (i.e. the vegetation was spatially structured). We defined three plant communities that differed along two main environmental gradients. The main gradient operates on a regional scale and is determined, from west to east, by a decrease in mean annual precipitation and an increase in the depth of the WT, soil pH, and EC. The secondary gradient operates on a site scale and is determined by topographic features inside the mallín and their influence on the hydrological regime (increasing moisture from the border towards the center). This second gradient allowed us to distinguish two plant communities, one of wet characteristics in the centers of the mallín, and another of mesic characteristics along the borders of the mallín.     

Disentangling the relative importance of climate,size and competition on tree growth in Iberian forests: implications for forest management under global change

Most large‐scale multispecies studies of tree growth have been conducted in tropical and cool temperate forests, whereas Mediterranean water‐limited ecosystems have received much less attention. This limits our understanding of how growth of coexisting tree species varies along environmental gradients in these forests, and the implications for species interactions and community assembly under current and future climatic conditions. Here, we quantify the absolute effect and relative importance of climate, tree size and competition as determinants of tree growth patterns in Iberian forests, and explore interspecific differences in the two components of competitive ability (competitive response and effect) along climatic and size gradients. Spatially explicit neighborhood models were developed to predict tree growth for the 15 most abundant Iberian tree species using permanent‐plot data from the Spanish Second and Third National Forest Inventory (IFN). Our neighborhood analyses showed a climatic and size effect on tree growth, but also revealed that competition from neighbors has a comparatively much larger impact on growth in Iberian forests. Moreover, the sensitivity to competition (i.e. competitive response) of target trees varied markedly along climatic gradients causing significant rank reversals in species performance, particularly under xeric conditions. We also found compelling evidence for strong species‐specific competitive effects in these forests. Altogether, these results constitute critical new information which not only furthers our understanding of important theoretical questions about the assembly of Mediterranean forests, but will also be of help in developing new guidelines for adapting forests in this climatic boundary to global change. If we consider the climatic gradients of this study as a surrogate for future climatic conditions, then we should expect absolute growth rates to decrease and sensitivity to competition to increase in most forests of the Iberian Peninsula (in all but the northern Atlantic forests), making these management considerations even more important in the future.     

Patterns in abundance and distribution of juvenile flounder, Platichthys flesus, in Minho estuary (NW Iberian Peninsula)

For 2 years, the flatfish community of Minho estuary (Northwest of Iberian Peninsula) was surveyed monthly with a beam trawl. Flounder, Platichthys flesus, was the most representative species contributing the most (74%) to the total flatfish abundance and was the only species occurring throughout the sampling period and along the entire salinity gradient. The majority of flounder sampled consisted of 0- and 1-year group individuals. In both years, a peak in abundance was recorded in late spring–summer months (from May to August) which coincided with seasonal settlement events. Higher densities of newly-settled flounder were obtained in the most upstream site, located in the tidal freshwater wetland, indicating that this area is an important nursery area. Of the environmental variables tested, temperature and salinity were the best predictors for 0-year group abundance explaining around 30% of the variability, whereas sediment was also important when considering all age groups together. This study supports the generally prevalent idea that estuaries act as nurseries for the early life stages of flounder, with particular relevance to the tidal freshwater estuarine wetlands.     

A field test of the stress‐gradient hypothesis along an aridity gradient

Aims: The stress‐gradient hypothesis (SGH) predicts how plant interactions change along environmental stress gradients. We tested the SGH in an aridity gradient, where support for the hypothesis and the specific shape of its response curve is controversial. Location: Almería, Cáceres and Coimbra, three sites in the Iberian Peninsula that encompass the most arid and wet habitats in the distribution range of a nurse shrub species –Retama sphaerocarpa L. (Boiss) – in Europe. Methods: We analysed the effect of Retama on its understorey plant community and measured the biomass and species richness beneath Retama and in gaps. We estimated the frequency (changes in species richness), importance and intensity of the Retama effects, and derived the severity–interaction relationship pattern, analysing how these interaction indices changed along this aridity gradient. Results and conclusions: The intensity and frequency of facilitation by Retama increased monotonically with increasing environmental severity, and the importance tended to have a similar pattern, overall supporting the SGH. Our data did not support predictions from recent revisions of the SGH, which may not apply to whole plant communities like those studied here or when interactions are highly asymmetrical. Facilitation by Retama influenced community composition and species richness to the point that a significant fraction of species found at the most arid end of the gradient were only able to survive beneath the nurse shrub, whereas some of these species were able to thrive in gaps at more mesic sites, highlighting the ecological relevance of facilitation by nurse species in mediterranean environments, especially in the driest sites.     

Biogeography of Iberian freshwater fishes revisited: the roles of historical versus contemporary constraints

Aim The question of how much of the shared geographical distribution of biota is due to environmental vs. historical constraints remains unanswered. The aim of this paper is to disentangle the contribution of historical vs. contemporary factors to the distribution of freshwater fish species. In addition, it illustrates how quantifying the contribution of each type of factor improves the classification of biogeographical provinces. Location Iberian Peninsula, south‐western Europe (c. 581,000 km²). Methods We used the most comprehensive data on native fish distributions for the Iberian Peninsula, compiled from Portuguese and Spanish sources on a 20‐km grid‐cell resolution. Overall, 58 species were analysed after being categorized into three groups according to their ability to disperse through saltwater: (1) species strictly intolerant of saltwater (primary species); (2) species partially tolerant of saltwater, making limited incursions into saltwaters (secondary species); and (3) saltwater‐tolerant species that migrate back and forth from sea to freshwaters or have invaded freshwaters recently (peripheral species). Distance‐based multivariate analyses were used to test the role of historical (basin formation) vs. contemporary environmental (climate) conditions in explaining current patterns of native fish assemblage composition. Cluster analyses were performed to explore species co‐occurrence patterns and redefine biogeographical provinces based on the distributions of fishes. Results River basin boundaries were better at segregating species composition for all species groups than contemporary climate variables. This historical signal was especially evident for primary and secondary freshwater fishes. Eleven biogeographical provinces were delineated. Basins flowing to the Atlantic Ocean north of the Tagus Basin and those flowing to the Mediterranean Sea north of the Mijares Basin were the most dissimilar group. Primary and secondary freshwater species had higher province fidelity than peripheral species. Main conclusions The results support the hypothesis that historical factors exert greater constraints on native freshwater fish assemblages in the Iberian Peninsula than do current environmental factors. After examining patterns of assemblage variation across space, as evidenced by the biogeographical provinces, we discuss the likely dispersal and speciation events that underlie these patterns.     

Spatial Distribution of Dorylaimid and Mononchid Nematodes from the Southeast Iberian Peninsula: Environmental Characterization of Chorotypes

The aim of this study was to determine the incidence of 18 environmental variables in the spatial distribution of 30 chorotypes (species groups with significantly similar distribution patterns) of dorylaimid and mononchid nematodes by means of logistic regression in a natural area in the southeastern Iberian Peninsula. Six variables (elevation, color chroma, clay content, nitrogen content, CaCO₃, and plant community associated) were the most important environmental factors that helped explain the distribution of chorotypes. The distribution of most chorotypes was characterized by some (one to three) environmental variables; only two chorotypes were characterized by five or more variables, and four have not been characterized.     

Habitat quality affects the condition of Barbus sclateri in Mediterranean semi-arid streams

We investigated the site level relationships between fish condition and environmental variables in Sclater's barbel, Barbus sclateri, from semi-arid streams in the south-eastern Iberian Peninsula. These freshwater ecosystems are characterized by strong seasonal fluctuations in flow levels (droughts and floods) as a consequence of their irregular hydrological regime on both an annual and pluriannual scale. We analysed the relationships between 11 ecosystem variables [conductivity, oxygen concentration, water temperature, pH, seasonal water flow, dominant substrate, submerged vegetation, QBR (Riparian Ecosystems Quality Index), BMWP' (Spanish version of the Biological Monitoring Working Party), fish refuge index and fish density] and fish condition by comparing mass–length relationships of nine populations of Sclater's barbel located in five sub-basins with different environmental conditions. Fish condition differed between the populations studied and was mainly dependent on the ecological variables related with water flow and, consequently, the physical structure of the streams, which is directly related to substrate and fish refuge.     

Effectiveness of a large reserve network in protecting freshwater biodiversity: a test for the Iberian Peninsula

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Comparative study of the hyperbenthos of three European estuaries

The hyperbenthic fauna of the subtidal channels of the Eems (N. Netherlands), Westerschelde (S.W. Netherlands), and Gironde (S.W. France) estuaries was sampled within a 15-day period in summer 1991. In each estuary, quantitative samples were taken at regularly spaced stations covering the entire salinity gradient from marine conditions at the mouth to nearly freshwater conditions upstream. The diversity of the samples and the distribution of the species along the main estuarine gradients were assessed. Hyperbenthic communities were identified using different multivariate statistical techniques. The species composition and the density and biomass of the dominant species of each community were compared among communities.     

Breeding Habitat Selection of Salamandra salamandra (Linnaeus, 1758) in the Most Arid Zone of its European Distribution Range: Application to Conservation Management

The influence of environmental variables on the selection of a water body as breeding habitat by Salamandra salamandra was studied in an arid zone located in the southwestern part of its distribution range. From November 2002 to October 2003, 50 water bodies were monitored in the south east of the Iberian Peninsula. Environmental data were submitted to a stepwise logistic regression analysis at macrohabitat, water body typology and microhabitat scales in order to establish the main factors influencing the use of a given water body as breeding habitat by this species. A significant degree of dependence between the reproduction of Salamandra salamandra and environmental variables was observed at all of these levels. These results should be taken into account when populations of this species are subjected to management and/or recovery programmes in arid areas.     

Environmental aridity is associated with cytotype segregation and polyploidy occurrence in Brachypodium distachyon (Poaceae)

? The ecological and adaptive significance of plant polyploidization is not well understood and no clear pattern of association between polyploid frequency and environment has emerged. Climatic factors are expected to predict cytotype distribution. However, the relationship among climate, cytotype distribution and variation of abiotic stress tolerance traits has rarely been examined. ? Here, we use flow cytometry and root-tip squashes to examine the cytotype distribution in the temperate annual grass Brachypodium distachyon in 57 natural populations distributed across an aridity gradient in the Iberian Peninsula. We further investigate the link between environmental aridity, ploidy, and variation of drought tolerance and drought avoidance (flowering time) traits. ? Distribution of diploids (2n = 10) and allotetraploids (2n = 30) in this species is geographically structured throughout its range in the Iberian Peninsula, and is associated with aridity gradients. Importantly, after controlling for geographic and altitudinal effects, the link between aridity and polyploidization occurrence persisted. Water-use efficiency varied between ploidy levels, with tetraploids being more efficient in the use of water than diploids under water-restricted growing conditions. ? Our results indicate that aridity is an important predictor of polyploid occurrence in B. distachyon, suggesting a possible adaptive origin of the cytotype segregation.