Miguel Clavero

Global ecological impacts of invasive species in aquatic ecosystems

The introduction of invasive species, which often differ functionally from the components of the recipient community, generates ecological impacts that propagate along the food web. This review aims to determine how consistent the impacts of aquatic invasions are across taxa and habitats. To that end, we present a global meta-analysis from 151 publications (733 cases), covering a wide range of invaders (primary producers, filter collectors, omnivores and predators), resident aquatic community components (macrophytes, phytoplankton, zooplankton, benthic invertebrates and fish) and habitats (rivers, lakes and estuaries). Our synthesis suggests a strong negative influence of invasive species on the abundance of aquatic communities, particularly macrophytes, zooplankton and fish. In contrast, there was no general evidence for a decrease in species diversity in invaded habitats, suggesting a time lag between rapid abundance changes and local extinctions. Invaded habitats showed increased water turbidity, nitrogen and organic matter concentration, which are related to the capacity of invaders to transform habitats and increase eutrophication. The expansion of invasive macrophytes caused the largest decrease in fish abundance, the filtering activity of filter collectors depleted planktonic communities, omnivores (including both facultative and obligate herbivores) were responsible for the greatest decline in macrophyte abundance, and benthic invertebrates were most negatively affected by the introduction of new predators. These impacts were relatively consistent across habitats and experimental approaches. Based on our results, we propose a framework of positive and negative links between invasive species at four trophic positions and the five different components of recipient communities. This framework incorporates both direct biotic interactions (predation, competition, grazing) and indirect changes to the water physicochemical conditions mediated by invaders (habitat alteration). Considering the strong trophic links that characterize aquatic ecosystems, this framework is relevant to anticipate the far-reaching consequences of biological invasions on the structure and functionality of aquatic ecosystems.

Climate Change or Land Use Dynamics: Do We Know What Climate Change Indicators Indicate?

Different components of global change can have interacting effects on biodiversity and this may influence our ability to detect the specific consequences of climate change through biodiversity indicators. Here, we analyze whether climate change indicators can be affected by land use dynamics that are not directly determined by climate change. To this aim, we analyzed three community-level indicators of climate change impacts that are based on the optimal thermal environment and average latitude of the distribution of bird species present at local communities. We used multiple regression models to relate the variation in climate change indicators to: i) environmental temperature; and ii) three landscape gradients reflecting important current land use change processes (land abandonment, fire impacts and urbanization), all of them having forest areas at their positive extremes. We found that, with few exceptions, landscape gradients determined the figures of climate change indicators as strongly as temperature. Bird communities in forest habitats had colder-dwelling bird species with more northern distributions than farmland, burnt or urban areas. Our results show that land use changes can reverse, hide or exacerbate our perception of climate change impacts when measured through community-level climate change indicators. We stress the need of an explicit incorporation of the interactions between climate change and land use dynamics to understand what are current climate change indicators indicating and be able to isolate real climate change impacts.

Threatening processes and conservation management of endemic freshwater fish in the Mediterranean basin: A review

Mediterranean endemic freshwater fish are among the most threatened biota in the world. The Mediterranean basin has experienced substantial reductions in precipitation and water availability, which will worsen with climate change. Current water policy is directed to increase water-supply demands, especially for agriculture, and not to improve water-use efficiency and implement integrated and sustainable water management. Illegal extractions are common, exacerbating problems for important protected areas. Management is needed to mitigate the conflicts between environmental water and human demand, and ensure availability of water to maintain ecological processes and Mediterranean freshwater biodiversity. Water availability is not the only threat, although it is exacerbated by pollution and invasive species. The uneven spatial distribution of threats across the Mediterranean basin requires different strategies to conserve freshwater biodiversity. Implementation of multi-national laws (e.g. Water Framework Directive in the European Union) will help future management of freshwater ecosystems. Management actions must be planned at whole-catchment scales, with collaboration among different countries and water-management authorities. The current reserve area is small compared with other areas in the world and driven by terrestrial interests, and should be evaluated for its effectiveness to protect the Mediterranean freshwater biodiversity.

Reservoirs promote the taxonomic homogenization of fish communities within river basins

Most studies analyzing patterns in biotic homogenization of fish communities have used large-scale approaches, while the community-level effects of species introductions and local extinctions within river basins have been sparsely analyzed. In this article, we examine patterns in freshwater fish α- and β-diversity in relation to the presence of reservoirs in a Mediterranean river (Guadiana river; Iberian Peninsula). We used fish samples from 182 river localities and 59 reservoir ones to address two main questions: (i) do reservoirs favor the establishment of invasive fish species?; and (ii) do reservoirs bear taxonomically homogenized fish communities? Although total species richness was not different between rivers and reservoirs, the latter had more invasive species and less native ones. Fish species found in reservoirs tended to be larger ones, but invasive species of any size showed higher preferences for reservoirs. Native species that were rare or absent in reservoirs were those that showed higher sensitivity to invasive species in rivers. Reservoir fish communities were taxonomically homogenized in relation to river ones, both when considering all fish species and using only natives or only invasive ones. Our results suggest that invasive species occupying reservoirs constitute an ecological filter excluding most native species from such systems. Invasive species in the study area are often widely introduced elsewhere, while native species found in reservoirs are congeneric and ecologically similar to those found in other Iberian studies. Thus, we conclude that reservoirs promote taxonomic homogenization at multiple spatial scales, while could also be promoting the functional homogenization of Iberian fish communities.

Bird community specialization, bird conservation and disturbance: the role of wildfires

1. Although niche theory predicts that disturbance should favour generalist species, the community-level implications of this pattern have been sparsely analysed. Here, we test the hypothesis that disturbance favours generalist species within communities, analysing effects of wildfires in bird communities in a Mediterranean climate area as a case study. 2. We use bird occurrence data in more than 500 1 × 1 km squares forming a gradient running from forest to completely burnt areas. The level of specialization of bird communities was estimated by means of three complementary species specialization indices, calculated for different landscape gradients and averaged at the community level (i.e. 1 × 1 km squares), and mean species rarity. 3. We also calculated mean habitat preferences along landscape gradients, as well as an index of conservation value and total species richness. 4. Different estimators of bird community specialization varied in contrasting fashion along the wildfire disturbance gradient, and thus we conclude that it is not justified to expect unique community responses to the sharp variations in habitat characteristics brought by wildfire disturbances. 5. Burnt areas tended to have rarer and urban-avoider bird species, whereas unburnt forests tended to have larger proportions of forest specialist species. 6. The mean conservation value of communities clearly increased towards the burnt extreme of the wildfire disturbance gradient, while this had a negligible effect on species richness. 7. Wildfires seem to play an important role for the maintenance of open-habitat, urban-avoider bird populations in Mediterranean landscapes and also to benefit a set of bird species of unfavourable European conservation status. 8. In this context, it cannot be unambiguously concluded that fire disturbance, even in a context in which fires are greatly favoured by human-related activities, leads to more functionally simplified communities dominated by generalist species.

Potential impact of an exotic mammal on rocky intertidal communities of northwestern Spain

Being the interface of sea and land, the coast can be invaded by introduced species coming from either of these two worlds. Recent reviews of coastal invasions emphasize the human-mediated transport of non-indigenous marine plants and invertebrates, forgetting the potential role of invaders of terrestrial origin. By studying the diet of the introduced American mink (Mustela vison) on a rocky shore of southwestern Europe, we draw attention to the potential impact on intertidal communities of exotic species coming from inland. We analysed 199 mink faeces collected in August 1997 and August 1999 in Baiona, a coastal and urban area of northern Spain recently invaded by minks. The diet of the species was based almost exclusively on crabs (45.4% of individual prey) and fish (53.3%). Most crabs were marbled crabs (Pachygrapsus marmoratus) and most fish were adult blennies (Coryphoblennius galerita and Lipophrys pholis). Given its energy requirements (about 1250 kJ/day), a single mink will consume during the month of August approximately 945 blennies and 496 crabs. Although we lack accurate data on mink abundance, a cautious estimation (4 mink/km before dispersal), supported by field observations, suggests that predation in August may reach 3780 blennies and 1984 crabs per km of shoreline. This predation pressure could affect the numbers of blennies and (less probably) crabs, indirectly benefiting the populations of their prey, that is, sessile invertebrates and snails. More field research is needed, but our results suggest that an exotic non-marine top predator such as the American mink could affect intertidal communities in Eurasia.

Territory characteristics and coexistence with heterospecifics in the Dartford warbler Sylvia undata across a habitat gradient

The study of successional gradients may help to understand the relative influence of habitat structure and competition on territory characteristics. Here, we evaluate the effects of vegetation cover, conspecific and heterospecific densities, and distance to the nearest neighbor on territory size, shape, and overlap in insectivorous birds. We studied these effects along a gradient of postfire habitat regeneration in which foliage cover and densities of focal species varied several-fold. We delineate 197 territories (minimum convex polygons) of the shrub-dwelling Dartford warbler (Sylvia undata) and 255 of the syntopic Sardinian (Sylvia melanocephala), subalpine (Sylvia cantillans) and melodious (Hippolais polyglotta) warblers at three plots in NE Catalonia (Spain and France) in 1987–2005. After accounting for the effect of the number of locations used to delineate polygons, generalized linear mixed models (GLMM) showed a reduction in territory area of the Dartford warbler as conspecific density increased and distance to nearest neighbor decreased, in accordance with the contender pressure hypothesis for territory size regulation. Heterospecific density was not included in the final model of territory size and the effect of habitat structure was marginal. Territory roundness was positively correlated with its size and with conspecific density, probably in relation to energetic constraints, and negatively with heterospecific density. Territorial exclusion was almost complete among Dartford warblers, whereas interspecific territory overlap was extensive and tended to increase with heterospecific density and with structural diversity along the gradient. Our results support the hypothesis that Mediterranean warbler coexistence derives from ecological segregation and not from interspecific territoriality.

Population and microhabitat effects of interspecific interactions on the endangered Andalusian toothcarp (Aphanius baeticus)

The Andalusian toothcarp, Aphanius baeticus, is a critically endangered cyprinodontid species, with only nine known extant populations. Although not yet studied in the field, the distribution and abundance of Andalusian toothcarp are thought to be strongly influenced by interspecific interactions. We analysed the abundance and microhabitat use of Andalusian toothcarp in two water courses, one in which several other fish species occurred (sympatric site) and one hypersaline stream in which toothcarp was the only species present (allopatric site). Fish were sampled using plastic minnow traps and results were analysed separately for three size categories. Toothcarps were clearly more abundant in the allopatric population than in the sympatric one, though the difference was less apparent in the smallest size category. In coexistence with other species, toothcarp occupied shallower microhabitats, but in both sites in the absence of shelter fish selected deeper positions than in its absence. While in the sympatric site sheltered microhabitats were used predominately by small individuals, in the allopatric ones they were used by larger ones. Observed patterns strongly suggest that predation is the main mechanism involved in the differences in abundance and microhabitat use between sites. Our results confirm that the presence or absence of coexisting species is an important habitat feature for Andalusian toothcarp populations.

Historical data to plan the recovery of the European eel

Summary 1. Long-term perspectives are critical to understand contemporary ecological systems. However, historical data on the distribution of biodiversity have only rarely been used in applied environmental sciences. 2. Here, we use historical sources to reconstruct the historical range of the European eel, a critically endangered species. We then use this baseline range to set range targets for the recovery of the European eel, as opposed to the abundance-based targets established by the European Union, which are constrained by the poor information on pre-collapse stocks. 3. We collected over 10 000 historical freshwater fish records from Spain in the 19th and 16th centuries, as well as over 25 000 records from the global biodiversity information facility (GBIF) to characterize historical and current European eel distribution in the Iberian Peninsula. We converted fish records into an eel presence–absence data set using subcatchment as spatial unit of analysis and modelled eel distribution in the different historical periods. 4. The eel was historically widespread throughout the Iberian Peninsula, but it has lost over 80% of its original range, mainly due to river fragmentation by dams. Distribution models applied to 16th- and 19th-century data showed a high agreement, supporting the use of the 19th-century estimated distribution as a baseline range. We identified the number and identity of dams that should be made passable for accomplishing specific range recovery targets, for example showing that acting upon 20 dams would make available 60% of the baseline eel range. 5. Synthesis and applications. This work exemplifies how insights gained from historical ecology can support and guide present-day management of migratory fishes. Similar approaches could be developed throughout Europe to plan the recovery of the eel, since there are large amounts of historical eel records. Historical baseline ranges for the eel range should be incorporated into the European Union legal mandates aimed at the recovery of the species.

Using historical accounts to set conservation baselines: the case of Lynx species in Spain

The knowledge of the historical range of organisms is necessary to understand distribution dynamics and their drivers as well as to set reference conditions and conservation goals. We reviewed written sources documenting the presence of lynxes in Spain between the sixteenth and nineteenth centuries, trying to infer whether Lynx records referred to the Iberian (Lynx pardinus) or the Eurasian (Lynx lynx) species. We compiled 151 spatially specific, non-redundant Lynx records, dating between 1572 and 1897. Records appeared split in two neatly separated areas, North and South. Arguably, all historical records from the South area correspond to Iberian lynx, since almost all of them fall within the estimated range of the species in the 1950s. This suggests that the core distribution area of the species remained relatively stable for centuries, until the second half of the twentieth century. The reported range in the 1950s is thus an appropriate scenario to set conservation goals for this critically endangered species. The North area extended from Galicia and northern Portugal to the Mediterranean, mainly occupying the Atlantic-climate area of northern Iberia. Reported direct measurements and the accumulation of indirect evidences suggest that most, if not all, northern Lynx records refer to the Eurasian lynx, which had been previously detected in archaeological sites of the area. This species was apparently present in the Iberian Peninsula until the early-nineteenth century. Our results provide an objective, baseline distribution of Lynx species in Spain useful to guide conservation efforts both at the Iberian and European levels.