Núria Bonada

Macroinvertebrate community structure and biological traits related to flow permanence in a Mediterranean river network

In Mediterranean climate areas, the great seasonal variability in temperature and rainfall is considered to be an evolutionary pressure that constrains plant and animal communities and their biological traits. Droughts alter habitat availability (changes of flow alter riffle-pool sequences), although habitat characteristics may also exacerbate drought to some extent. Using a simple quantitative index based on the proportion of conglomerate bedrock versus gravel and cobbles, pools versus riffles and winter versus summer flow, we show how habitat characteristics (in terms of substratum and flow) may influence the permanency of a stream site and how flow permanence constrains macroinvertebrate community structure and biological traits. Annual and seasonal macroinvertebrate richness, and the EPT (Ephemeroptera, Plecoptera and Trichoptera) and OCH (Odonata, Coleoptera and Hemiptera) metrics differed between permanent, intermittent and ephemeral sites, but not between permanent and intermittent sites. In contrast, distinct biological traits were observed in the three flow categories, although permanent sites presented few significant traits which was attributed to the stability of the habitat. Intermittent sites were dominated by taxa with pool-like strategies, while ephemeral sites were characterized by fauna with life-history adaptations to floods and droughts. In contrast to most traits (e.g., dissemination, reproduction, substrate relation), which were more constrained by local flow and substrate characteristics, life-cycle characteristics did not differ significantly among flow categories. This pattern can be explained by the features of the Mediterranean climate and particularly its high seasonal predictability, which serves as a large-scale filter of life-cycle traits, independently of local hydromorphological characteristics. Our findings indicate that drought is related to habitat characteristics and that local habitat variability favours organisms with certain traits, while other traits are independent of habitat variability on this scale and are probably affected by other large-scale habitat characteristics.

Benthic macroinvertebrate assemblages and macrohabitat connectivity in Mediterranean-climate streams of northern California

Abstract Drought leads to a loss of longitudinal and lateral hydrologic connectivity, which causes direct or indirect changes in stream ecosystem properties. Changes in macrohabitat availability from a riffle–pool sequence to isolated pools are among the most conspicuous consequences of connectivity loss. Macroinvertebrate assemblages were compared among 3 distinct stream macrohabitats (riffles [R], pools connected to riffles [Pc], disconnected pools [Pd]) of 19 Mediterranean-climate sites in northern California to examine the influence of loss of habitat resulting from drought disturbance. At the time of sampling, 10 sites were perennial and included R and Pc macrohabitats, whereas 9 sites were intermittent and included only Pd macrohabitats. Taxa richness was more variable in Pd, and taxa richness was significantly lower in Pd than in Pc but not R. These results suggested a decline in richness between Pc and Pd that might be associated with loss of connectivity. Lower Ephemeroptera, Plecoptera, and Trichoptera (EPT) richness relative to Odonata, Coleoptera, and Heteroptera (OCH) richness was observed for Pd than R and Pc macrohabitats. Family composition was more similar between R and Pc than between R or Pc and Pd macrohabitats. This similarity may be associated with greater connectivity between R and Pc macrohabitats. Correspondence analysis indicated that macroinvertebrate composition changed along a gradient from R to Pc and Pd that was related to a perennial–intermittent gradient across sites. High variability among macroinvertebrate assemblages in Pd could have been related to variability in the duration of intermittency. In cluster analysis, macroinvertebrate assemblages were grouped by macrohabitat first and then by site, suggesting that the macrohabitat filter had a greater influence on macroinvertebrate assemblages than did local site characteristics. Few taxa were found exclusively in Pc, and this macrohabitat shared numerous taxa with R and Pd, indicating that Pc may act as a bridge between R and Pd during drought. Drought is regarded as a ramp disturbance, but our results suggest that the response of macroinvertebrate assemblages to the loss of hydrological connectivity among macrohabitats is gradual, at least in Mediterranean-climate streams where drying is gradual. However, the changes may be more dramatic in arid and semiarid streams or in Mediterranean-climate streams if drying is rapid.

Biomonitoring of human impacts in freshwater ecosystems:the good, the bad and the ugly

It is critical that the impacts of environmental stressors on natural systems are detected, monitored and assessed accurately in order to legislate effectively and to protect and restore ecosystems. Biomonitoring underpins much of modern resource management, especially in fresh waters, and has received significant sums of money and research effort during its development. Despite this, the incorporation of science has not been effective and the management tools developed are sometimes inappropriate and poorly designed. Much biomonitoring has developed largely in isolation from general ecological theory, despite the fact that many of its fundamental principles ultimately stem from basic concepts, such as niche theory, the habitat template and the r–K continuum. Consequently, biomonitoring has not kept pace with scientific advances, which has compromised its ability to deal with emerging environmental stressors such as climate change and habitat degradation. A reconnection with its ecological roots and the incorporation of robust statistical frameworks are key to progress and meeting future challenges. The vast amount of information already collected represents a potentially valuable, and largely untapped, resource that could be used more effectively in protecting ecosystems and in advancing general ecology. Biomonitoring programmes have often accumulated valuable long-term data series, which could be useful outside the scope of the original aims. However, it is timely to assess critically existing biomonitoring approaches to help ensure future programmes operate within a sound scientific framework and cost-effectively. Investing a small proportion of available budgets to review effectiveness would pay considerable dividends. Increasing activity has been stimulated by new legislation that carries the threat of penalties for non-compliance with environmental targets, as is proposed, for example, in the EUs Water Framework Directive. If biomonitoring produces poor-quality data and has a weak scientific basis, it may lead either to unjustified burdens placed on the users of water resources, or to undetected environmental damage. We present some examples of good practice and suggest new ways to strengthen the scientific rigour that underpins biomonitoring programmes, as well as highlighting potentially rewarding new approaches and technologies that could complement existing methods.

Small but mighty: headwaters are vital to stream network biodiversity at two levels of organization

Abstract Headwaters (stream orders 1–2) traditionally have been considered depauperate compared to mid-order streams (orders 3–4)—a conclusion that arises from a perception of streams as linear systems and emphasizes change in average &agr; (local) diversity along streams. We hypothesized an opposite pattern for &bgr; (among-site) diversity and suggest that headwaters might account for a large degree of basin-scale biodiversity if considered within the more realistic framework of streams as branching networks. We assembled pre-existing biodiversity data from across the globe to test this hypothesis broadly at the population-genetic (mitochondrial haplotype diversity within species) and community (species/taxonomic diversity) levels, with a focus on macroinvertebrates. We standardized 18 (9 headwater and 9 mid-order) population-genetic and 16 (10 headwater and 6 mid-order) community-level ecoregional data sets from 5 global ecozones for robust comparisons of &bgr;-diversity estimates between the 2 stream-size categories. At the population-genetic level, we applied measures of among-site variation commonly used at both population-genetic (FST and &PHgr;ST) and community (Sørensens dissimilarity with both presence/absence and abundance data) levels and developed a novel strategy to compare expected rates of loss of &ggr; (regional) diversity as individual sites are eliminated sequentially from regions. At the community level, we limited analyses to Sørensens presence/absence measures. We found that Sørensens dissimilarity was significantly greater among headwaters than among mid-order streams at both population-genetic and community levels. We also showed that individual headwater reaches accounted for greater proportions of genetic &ggr; diversity than did mid-order reaches. However, neither FST nor &PHgr;ST was significantly different between stream-size categories. These measures, which have been used traditionally for comparisons of population-genetic variation, measure proportions of total variation rather than solely among-site variation (i.e., they also are influenced by within-site variation). In contrast, Sørensens dissimilarity measures only among-site variation and, therefore, is presumably more useful for reflecting general &bgr; diversity. Overall results suggest that, on average, headwaters probably contribute disproportionately to biodiversity at the network scale. This finding demands a shift in thinking about the biodiversity contributions of small headwaters and has strong conservation implications for imperiled headwater streams around the world.

Mediterranean-climate streams and rivers: geographically separated but ecologically comparable freshwater systems

Streams and rivers in mediterranean-climate regions (med-rivers in med-regions) are ecologically unique, with flow regimes reflecting precipitation patterns. Although timing of drying and flooding is predictable, seasonal and annual intensity of these events is not. Sequential flooding and drying, coupled with anthropogenic influences make these med-rivers among the most stressed riverine habitat worldwide. Med-rivers are hotspots for biodiversity in all med-regions. Species in med-rivers require different, often opposing adaptive mechanisms to survive drought and flood conditions or recover from them. Thus, metacommunities undergo seasonal differences, reflecting cycles of river fragmentation and connectivity, which also affect ecosystem functioning. River conservation and management is challenging, and trade-offs between environmental and human uses are complex, especially under future climate change scenarios. This overview of a Special Issue on med-rivers synthesizes information presented in 21 articles covering the five med-regions worldwide: Mediterranean Basin, coastal California, central Chile, Cape region of South Africa, and southwest and southern Australia. Research programs to increase basic knowledge in less-developed med-regions should be prioritized to achieve increased abilities to better manage med-rivers.

Drivers and stressors of freshwater biodiversity patterns across different ecosystems and scales: a review

The present review with focus on the last decade (2000–2010) aims to (i) collecting the major hypotheses explaining freshwater biodiversity patterns, (ii) identifying the main stressors affecting freshwater biodiversity, and (iii) revealing information gaps regarding ecosystem types, organism groups, spatial and temporal scales to highlight research needs to better propose sound conservation measures. The comparative analysis addresses six organism groups ranging from microorganisms to fish in basins, rivers, lakes, wetlands, ponds and groundwater. Short-term studies at ecoregion and catchment scale focusing on invertebrates, macrophytes and fish in Palaearctic and Nearctic regions dominated. The most frequent hypotheses tested were the landscape filter concept, the species–area relationship, the metacommunity concept. Dominating natural drivers were area, heterogeneity and disturbance. Land use, eutrophication and habitat destruction were identified as most important stressors. Generally, freshwater biodiversity declined in response to these stressors in contrast to increasing biodiversity determined by natural drivers across all ecosystems. Preferred organism groups were fish and invertebrates, most frequently studied in rivers, in contrast to smaller organisms (e.g. bacteria) and, e.g. groundwater being underrepresented. Hypotheses originating from the last century are still tested in freshwater research, while novel concepts are either missing or untested. Protection of freshwater biodiversity is the ultimate challenge since it supports valuable ecosystems services ensuring perpetuation of mankind. For that, comprehensive large-scale studies with holistic approaches are urgently needed.

A comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity-based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within-basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.

Ecological and historical filters constraining spatial caddisfly distribution in Mediterranean rivers

Summary 1. Contemporary species distributions are determined by a mixture of ecological and historical filters acting on several spatial and temporal scales. Mediterranean climate areas are one of the worlds biodiversity hotspots with a high level of endemicity, which is linked to complex ecological and historical factors. 2. This paper explores the ecological and historical factors constraining the distribution of caddisfly species on a large regional scale. A total of 69 taxa were collected from 140 sampling sites in 10 Iberian Mediterranean river basins. Approximately 74% of taxa can be considered rare, with the southern basins (the Baetic–Riffian region) having greater endemicity. The greatest richness, involving a mixture of northern and southern species, was found in the transitional area between the Baetic–Riffian region and the Hesperic Massif. 3. The historical processes occurring during the Tertiary (i.e. the junction of the Eurasian and African plates) explained 3.1% of species distribution, whereas ecological factors accounted for 20.7%. Only 0.3% was explained by the interaction of history and ecology. A set of multi‐scale ecological variables (i.e. basin, reach and bedform characteristics) defined five river types with specific caddisfly assemblages. The commonest caddisfly species accounted for the regional distribution pattern, while rare taxa contributed to the explanation of subtle patterns not shown by common species. 4. Despite the importance of historical factors for biogeography and the large scale used in our study, ecological variables better explained caddisfly distribution. This may be explained by the length of time since the historical process we are considering, the high dispersion and colonisation capacity of many caddisfly species, and the strong environmental gradient in the area. Because of the historical and environmental complexity of Mediterranean areas, rare taxa should be included in ecological studies so that the singularity of these ecosystems is not missed.

Vulnerability of stream biota to climate change in mediterranean climate regions: a synthesis of ecological responses and conservation challenges

Freshwater species worldwide are experiencing dramatic declines partly attributable to ongoing climate change. It is expected that the future effects of climate change could be particularly severe in mediterranean climate (med-) regions, which host many endemic species already under great stress from the high level of human development. In this article, we review the climate and climate-induced changes in streams of med-regions and the responses of stream biota, focusing on both observed and anticipated ecological responses. We also discuss current knowledge gaps and conservation challenges. Expected climate alterations have already been observed in the last decades, and include: increased annual average air temperatures; decreased annual average precipitation; hydrologic alterations; and an increase in frequency, intensity and duration of extreme events, such as floods, droughts and fires. Recent observations, which are concordant with forecasts built, show stream biota of med-regions when facing climate changes tend to be displaced towards higher elevations and upper latitudes, communities tend to change their composition and homogenize, while some life-history traits seem to provide biota with resilience and resistance to adapt to the new conditions (as being short-lived, small, and resistant to low streamflow and desiccation). Nevertheless, such responses may be insufficient to cope with current and future environmental changes. Accurate forecasts of biotic changes and possible adaptations are difficult to obtain in med-regions mainly because of the difficulty of distinguishing disturbances due to natural variability from the effects of climate change, particularly regarding hydrology. Long-term studies are needed to disentangle such variability and improve knowledge regarding the ecological responses and the detection of early warning signals to climate change. Investments should focus on taxa beyond fish and macroinvertebrates, and in covering the less studied regions of Chile and South Africa. Scientists, policy makers and water managers must be involved in the climate change dialogue because the freshwater conservation concerns are huge.

Biological attributes discriminating invasive from native European stream macroinvertebrates

Rising economic and ecological costs caused by invasive organisms revived research on biological attributes associated with invasiveness, focussing on the question: do invasive taxa have biological attributes favouring (1) propagule pressure; (2) dispersal; and (3) establishment and population growth? Using a literature-derived database on 312 stream macroinvertebrate genera occurring at 527 least human-impacted European sites, we quantitatively examined this question. Compared with native genera, genera with invasive invertebrate species (1) tended to reproduce more frequently and to have higher abundances (i.e. higher propagule pressure); (2) had no particular resistance stages to survive during dispersal; and (3) had significantly more ovoviviparity (enabling colonization by a single individual that typically releases viable offspring), larger size and longer life (providing resistance against mortality), food and feeding habits exploiting food resources in streams more effectively, and tended to be more dominant in their communities (all favouring establishment and population growth). Repeating these analyses excluding “native” flying insects (i.e. genera that presumably invaded from glacial refuges unnoticed by biologists), fewer biological attributes were significantly associated with invasiveness. For both data sets (all genera or insects excluded), their affinity to few biological traits (e.g. ovoviviparity, gill respiration) assigned the same 13 of the 19 invasive genera to the top 19 ranks on a gradient of potential invasiveness, together with native genera that risk to become invasive (e.g. Pisidium, Unio), but also with one endangered native genus (Margaritifera). Overall, our data support the idea that invasiveness can be predicted using biological attributes.