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.

Freshwater biodiversity and conservation in mediterranean climate streams of Chile.

In Chile, mediterranean climate conditions only occur in the Central Zone (ChMZ). Despite its small area, this mediterranean climate region (med-region) has been recognised as a hotspot for biodiversity. However, in contrast to the rivers of other med-regions, the rivers in the ChMZ have been studied infrequently, and knowledge of their freshwater biodiversity is scarce and fragmented. We gathered information on the freshwater biodiversity of ChMZ, and present a review of the current knowledge of the principal floral and faunal groups. Existing knowledge indicates that the ChMZ has high levels of endemism, with many primitive species being of Gondwanan origin. Although detailed information is available on most floral groups, most faunal groups remain poorly known. In addition, numerous rivers in the ChMZ remain completely unexplored. Taxonomic specialists are scarce, and the information available on freshwater biodiversity has resulted from studies with objectives that did not directly address biodiversity issues. Research funding in this med-region has a strong applied character and is not focused on the knowledge of natural systems and their biodiversity. Species conservation policies are urgently required in this highly diverse med-region, which is also the most severely impacted and most populated region of the country.

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.

Identification and conservation of remnant genetic resources of brown trout in relict populations from Western Mediterranean streams

Brown trout is a cold-adapted freshwater species with restricted distribution to headwater streams in rivers of the South European peninsulas, where populations are highly vulnerable because Mediterranean regions are highly sensitive to the global climatic warming. Moreover, these populations are endangered due to the introgressive hybridization with cultured stocks. Individuals from six remnant populations in Western Mediterranean rivers were sequenced for the complete mitochondrial DNA control region and genotyped for 11 nuclear markers. Three different brown trout lineages were present in the studied region. Significant genetic divergence was observed among locations and a strong effect of genetic drift was suggested. An important stocking impact (close to 25%) was detected in the zone. Significant correlations between mitochondrial-based rates of hatchery introgression and water flow variation suggested a higher impact of stocked females in unstable habitats. In spite of hatchery introgression, all populations remained highly differentiated, suggesting that native genetic resources are still abundant. However, climatic predictions indicated that suitable habitats for the species in these rivers will be reduced and hence trout populations are highly endangered and vulnerable. Thus, management policies should take into account these predictions to design upstream refuge areas to protect remnant native trout in the region

Vegetative and reproductive morphology of Kallymenia patens (Kallymeniaceae, Rhodophyta) in the Mediterranean Sea

Reproductive morphology of the Mediterranean red alga Kallymenia patens is described for the first time, confirming its position in the genus. K. patens is characterized by a non-procarpic female reproductive apparatus, carpogonial branch systems consisting of supporting cells bearing both three-celled carpogonial branches and subsidiary cells that lack a hypogynous cell and carpogonium; fusion cells develop numerous connecting filaments, and tetrasporangia are scattered over the thallus and are probably cruciately divided. Old fertile spathulate specimens of K. patens are morphologically similar to K. spathulata, but they can be distinguished by the length of spathulated proliferations (up to 0.6 cm and 6 cm, respectively), the length of inner cortical cells (up to 70 and 30 μm, respectively), and the gonimoblast location (in proliferations from the perennial part of the blade and over all the thallus surface, respectively)

The morphology and reproductive structures of Mediterranean species of the genus Nemastoma J. Agardh, nom. cons. (Nemastomataceae, Nemastomatales): Nemastoma dichotomum and N. dumontioides

The morphology and reproductive structures of Mediterranean species of the genus Nemastoma J. Agardh, nom. cons. (Nemastomataceae, Nemastomatales): Nemastoma dichotomum and N. dumontioides

Geographic patterns of genetic variation in a broadly distributed marine vertebrate: new insights into loggerhead turtle stock structure from expanded mitochondrial DNA sequences

Previous genetic studies have demonstrated that natal homing shapes the stock structure of marine turtle nesting populations. However, widespread sharing of common haplotypes based on short segments of the mitochondrial control region often limits resolution of the demographic connectivity of populations. Recent studies employing longer control region sequences to resolve haplotype sharing have focused on regional assessments of genetic structure and phylogeography. Here we synthesize available control region sequences for loggerhead turtles from the Mediterranean Sea, Atlantic, and western Indian Ocean basins. These data represent six of the nine globally significant regional management units (RMUs) for the species and include novel sequence data from Brazil, Cape Verde, South Africa and Oman. Genetic tests of differentiation among 42 rookeries represented by short sequences (380 bp haplotypes from 3,486 samples) and 40 rookeries represented by long sequences (~800 bp haplotypes from 3,434 samples) supported the distinction of the six RMUs analyzed as well as recognition of at least 18 demographically independent management units (MUs) with respect to female natal homing. A total of 59 haplotypes were resolved. These haplotypes belonged to two highly divergent global lineages, with haplogroup I represented primarily by CC-A1, CC-A4, and CC-A11 variants and haplogroup II represented by CC-A2 and derived variants. Geographic distribution patterns of haplogroup II haplotypes and the nested position of CC-A11.6 from Oman among the Atlantic haplotypes invoke recent colonization of the Indian Ocean from the Atlantic for both global lineages. The haplotypes we confirmed for western Indian Ocean RMUs allow reinterpretation of previous mixed stock analysis and further suggest that contemporary migratory connectivity between the Indian and Atlantic Oceans occurs on a broader scale than previously hypothesized. This study represents a valuable model for conducting comprehensive international cooperative data management and research in marine ecology.

Flash flood evolution in North-Western Mediterranean

The present paper shows an in-depth analysis of the evolution of floods and precipitation in Catalonia for the period 1981-2010. In order to have homogeneous information, and having in mind that not gauge data was available for all the events, neither for all the rivers and stream flows, daily press from a specific newspaper has been systematically analysed for this period. Furthermore a comparison with a longer period starting in 1900 has been done. 219 flood events (mainly flash flood events) have been identified for the period of 30 years (375 starting in 1900), 79 of them were ordinary, 117 of them were extraordinary and 23 of them were catastrophic, being autumn and summer the seasons with the maxima values. 19% of the events caused a total of 110 casualties. 60% of them died when they tried to cross the street or the stream. Factors like the evolution of precipitation, population density and other socio-economical aspects have been considered. The trend analysis shows an increase of 1 flood/decade that probably has been mainly due to inter-annual and intra-annual changes in population density and in land-use and land-cover.

Spiders on a Hot Volcanic Roof: Colonisation Pathways and Phylogeography of the Canary Islands Endemic Trap-Door Spider Titanidiops canariensis (Araneae, Idiopidae)

Studies conducted on volcanic islands have greatly contributed to our current understanding of how organisms diversify. The Canary Islands archipelago, located northwest of the coast of northern Africa, harbours a large number of endemic taxa. Because of their low vagility, mygalomorph spiders are usually absent from oceanic islands. The spider Titanidiops canariensis, which inhabits the easternmost islands of the archipelago, constitutes an exception to this rule. Here, we use a multi-locus approach that combines three mitochondrial and four nuclear genes to investigate the origins and phylogeography of this remarkable trap-door spider. We provide a timeframe for the colonisation of the Canary Islands using two alternative approaches: concatenation and species tree inference in a Bayesian relaxed clock framework. Additionally, we investigate the existence of cryptic species on the islands by means of a Bayesian multi-locus species delimitation method. Our results indicate that T. canariensis colonised the Canary Islands once, most likely during the Miocene, although discrepancies between the timeframes from different approaches make the exact timing uncertain. A complex evolutionary history for the species in the archipelago is revealed, which involves two independent colonisations of Fuerteventura from the ancestral range of T. canariensis in northern Lanzarote and a possible back colonisation of southern Lanzarote. The data further corroborate a previously proposed volcanic refugium, highlighting the impact of the dynamic volcanic history of the island on the phylogeographic patterns of the endemic taxa. T. canariensis includes at least two different species, one inhabiting the Jandia peninsula and central Fuerteventura and one spanning from central Fuerteventura to Lanzarote. Our data suggest that the extant northern African Titanidiops lineages may have expanded to the region after the islands were colonised and, hence, are not the source of colonisation. In addition, T. maroccanus may harbour several cryptic species.

How are riparian plants distributed along the riverbank topographic gradient in Mediterranean rivers? Application to minimally altered river stretches in Southern Spain.

Species structure and composition in Mediterranean riparian forests are determined by hydrological features, longitudinal zonation, and riverbank topography. This study assesses the distribution of four native riparian plants along the riverbank topographic gradient in three river stretches in southern Spain, with special emphasis on the occupation of adult and young feet of each species. The studied stretches suffered minimal human disturbances, displayed semi-arid conditions, and had wide riparian areas to allow the development of the target species: black alder (Alnus glutinosa), salvia leaf willow (Salix salviifolia), narrow-leafed ash (Fraxinus angustifolia), and oleander (Nerium oleander). Thalweg height was used to define the riverbank topographic gradient. The results showed a preferential zone for black alder and salvia leaf willow in the range of 0-150 cm from the channel thalweg, with adult alders and willows being more common between 51 and 150 cm and young alders being more common under 50 cm. Conversely, narrow-leafed ash and oleander were much more frequent, and showed greater development, in the ranges of 151-200 cm and 201-250 cm, respectively, whereas the young feet of both species covered the entire topographic range. Adult feet of the four species were spatially segregated along the riverbank topographic gradient, indicating their differential ability to cope with water stress from the non-tolerant alders and willows to more tolerant narrow-leafed ash trees and oleanders. Young feet, however, showed a strategy more closely linked to the initial availability of colonisation sites within riparian areas to the dispersion strategy of each species and to the distribution of adult feet. In Mediterranean areas, where riparian management has traditionally faced great challenges, the incorporation of species preferences along riverbank gradients could improve the performance of restoration projects.

Modelling nitrogen and phosphorus loads in a Mediterranean river catchment (La Tordera, NE Spain)

Human activities have resulted in increased nutrient levels in many rivers all over Europe. Sustainable management of river basins demands an assessment of the causes and consequences of human alteration of nutrient flows, together with an evaluation of management options. In the context of an integrated and interdisciplinary environmental assessment (IEA) of nutrient flows, we present and discuss the application of the nutrient emission model MONERIS (MOdelling Nutrient Emissions into River Systems) to the Catalan river basin, La Tordera (north-east Spain), for the period 1996–2002. After a successful calibration and verification process (Nash-Sutcliffe efficiencies E=0.85 for phosphorus and E=0.86 for nitrogen), the application of the model MONERIS proved to be useful in estimating nutrient loads. Crucial for model calibration, in-stream retention was estimated to be about 50 % of nutrient emissions on an annual basis. Through this process, we identified the importance of point sources for phosphorus emissions (about 94% for 1996–2002), and diffuse sources, especially inputs via groundwater, for nitrogen emissions (about 31% for 1996–2002). Despite hurdles related to model structure, observed loads, and input data encountered during the modelling process, MONERIS provided a good representation of the major interannual and spatial patterns in nutrient emissions. An analysis of the model uncertainty and sensitivity to input data indicates that the model MONERIS, even in data-starved Mediterranean catchments, may be profitably used by water managers for evaluating quantitative nutrient emission scenarios for the purpose of managing river basins. As an example of scenario modelling, an analysis of the changes in nutrient emissions through two different future scenarios allowed the identification of a set of relevant measures to reduce nutrient loads.

Feeding ecology and trophic position of three sympatric demersal chondrichthyans in the Northwestern Mediterranean.

Understanding how marine predators interact is a scientific challenge. In marine ecosystems, segregation in feeding habits has been largely described as a common mechanism to allow the coexistence of several competing marine predators. However, little is known about the feeding ecology of most species of chondrichthyans, which play a pivotal role in the structure of marine food webs worldwide. In this study, we examined the trophic ecology of 3 relatively abundant chondrichthyans coexisting in the Mediterranean Sea: the blackmouth catshark Galeus melastomus , the velvet belly lanternshark Etmopterus spinax and the rabbit fish Chimaera monstrosa. To examine their trophic ecology and interspecific differences in food habits, we combined the analysis of stomach content and stable isotopes. Our results highlighted a trophic segregation between C. monstrosa and the other 2 species. G. melastomus showed a diet composed mainly of cephalopods, while E. spinax preyed mainly on shrimps and C. monstrosa on crabs. Interspecific differences in the trophic niche were likely due to different feeding capabilities and body size. Each species showed different isotopic niche space and trophic level. Specifically, C. monstrosa showed a higher trophic level than E. spinax and G. melastomus. The high trophic levels of the 3 species highlighted their important role as predators in the marine food web. Our results illustrate the utility of using complementary approaches that provide information about the feeding behaviour at short (stomach content) and long-term scales (stable isotopes), which could allow more efficient monitoring of marine food-web changes in the study area.

Crustacean and aquatic insect assemblages in the Mediterranean coastal ecosystems of Empord`a wetlands (NE Iberian peninsula)

Coastal wetlands are characterized by high biodiversity, which is one of the main criteria considered when establishing protection policies or when proposing adequate management actions. In this study, the crustacean and aquatic insect composition of the Empord`a wetlands is described. These two faunal groups contribute highly to the total biodiversity in these wetlands but are seldom considered when managing natural areas. A selection (84 sampling points) of all water body types present in the Empord`a wetlands were sampled monthly (surber and dip net with a 250 μm mesh). Sampling was carried out during 3 surveys (1991-92, 1996-97 and 1999-2000). A rich fauna of 125 crustacean and 295 aquatic insect taxa was identified. We characterized each water body type using the most abundant species and the relative species richness of the taxonomic groups. A classification of the water body types, according to similarity between inventories, groups the brackish and hyperhaline systems in one cluster and the various freshwater systems in another one. Among freshwater systems, lotic waters and freshwater wetlands have a high similarity, whereas rice fields and freshwater springs have a low similarity

Estimation of Water Circulation in a Mediterranean Salt Marsh and its Relationship with Flooding Causes

Variations in water volume in small depressions in Mediterranean salt marshes in Girona (Spain) are described and the potential causes for these variations analysed. Although the basins appear to be endorrheic, groundwater circulation is intense, as estimated from the difference between water volume observed and that expected from the balance precipitation / evaporation. The rate of variation in volume (VR = AV / VAt) may be used to estimate groundwater supply ('circulation'), since direct measurements of this parameter are impossible. Volume.conductivity figures can also be used to estimate the quantity of circulation, and to investigate the origin of water supplied to the system. The relationships between variations in the volume of water in the basins and the main causes of flooding are also analysed. Sea storms, rainfall levels and strong, dry northerly winds are suggested as the main causes of the variations in the volumes of basins. The relative importance assigned to these factors has changed, following the recent regulation of freshwater flows entering the system

Hydrology, light and the use of organic and inorganic materials as structuring factors of biological communities in Mediterranean streams

Hydrological disturbances, light availability and nutrients are the most relevant factors determining the structure of the biological communities in Mediterranean rivers. While some hydrological disturbances are able to induce catastrophic effects, which may cause a complete reset in physical and biological conditions, continued enrichment or changes in light availability are factors leading to the progressive shift in the communities of autotrophs and heterotrophs in the systems. Primary production in Mediterranean streams shows relevant seasonal changes which mainly follows the variations in light availability. In most forested streams, the algal community is shade-adapted. Nutrient enrichment (especially phosphorus) leads to marked increases in primary production, but this increase is not lineal and there is a saturation of algal biomass even in the most enriched systems. The heterotrophs (bacteria, fungi) are related to the pattern of DOC availability (which most depends on the seasonal discharge and leaf fall dynamics) and to the available substrata in the stream. It has been repeatedly observed that shorttime increases of extracellular enzyme activities are related to the accumulation of autochthonous (algal) and/or allochthonous (leaves) organic matter on the streambed during spring and summer, this being more remarkable in dry than in wetter years. Flow reduction favours detritus concentration in pools, and the subsequent increase in the density and biomass of the macroinvertebrate community. In Mediterranean streams collectors are accounting for the highest density and biomass, this being more remarkable in the least permanent systems, in accordance with the effect of floods on the organic matter availability. Nutrients, through the effect on the primary producers, also affect the trophic food web in the streams by favouring the predominance of grazers