1. Despite wide recognition that fish assemblages are influenced by factors operating over a range of spatial scales, little effort has been devoted to quantifying large‐scale variation and the multiscale dependencies of assemblage patterns and processes. This is particularly true for Mediterranean streams, where seasonally predictable drying‐up may lead to a strong association between assemblage attributes and large‐scale factors affecting the distribution of population sources and extinction likelihood. 2. The contribution of large‐scale factors to stream fish assemblage variation was quantified across a Mediterranean landscape, in south‐west Portugal. Fish abundance and species composition were estimated at 166 sites across third‐ to sixth‐order streams, in March–July 1998. Variance partitioning by redundancy analyses was used to analyse assemblage variation against three sets of predictor variables: environmental (catchment position, and geomorphic and hydrological factors), large‐scale spatial trends and neighbourhood effects. 3. Environmental variables and spatial trends accounted for 34.6% of the assemblage variation across the entire region, and for 36.6 and 57.8% within the two largest catchments (Mira and Seixe). Neighbourhood effects were analysed at the catchment scale, increasing the explained variation to 56.1% (Mira) and 70.7% (Seixe). 4. A prevailing environmental gradient was reflected in an increase in the abundance of all species and size‐classes in relation to catchment position, with more fish present in larger streams and in downstream reaches. Variables describing geomorphic and hydrological settings were less important in explaining assemblage variation. 5. Spatial trends always accounted for the smallest fraction of assemblage variation, and they were probably associated with historical barriers to fish dispersal. The strong neighbourhood effects may be related to spatially autocorrelated habitat conditions, but they are also a likely consequence of fish emigration/extinction and colonisation processes. 6. These results emphasise that a substantial proportion of fish assemblage variation in Mediterranean streams may be explained by large‐scale factors, irrespective of microhabitats and local biotic interactions. It is suggested that this pattern results to a large extent from the seasonal drying‐up, with the summer shortage of surface water limiting fish occurrence in headwaters, and consequently the key core areas for fish concentrating in larger streams and tributaries adjacent to large streams because of neighbourhood effects.
1. This study analysed changes occurring in Mediterranean stream fish assemblages over a sequence of dry years followed by a generally wet period . Variations in assemblage attributes were quantified at the basin and stream reach scales, and related to variables reflecting the occurrence of unusually dry or wet conditions. 2. Assemblage variability increased along with the resolution of analysis, with little changes in species richness, composition and rank abundances, but significant variation in individual species abundances. Fluctuations in relative abundances were significantly affected by variables reflecting the severity of summer droughts and the occurrence of rainy springs. These patterns were evident at the basin scale, while variability at individual stream reaches tended to be higher and less related to rainfall patterns. 3. At least three response guilds to rainfall variation could be identified: two of the four abundant and widespread species (chub and loach) declined following dry years, whereas the two other core species (nase and eel) declined after rainy spring; one scarce native species (stickleback) increased in dry years. 4. Except at the two most upstream reaches, the assemblages tended to recover quickly to previous configuration after the changes occurring during the sequence of dry years. 5. Temporal variability of local assemblages was concordant among reaches but did not follow any consistent spatial pattern. Instead, spatial patterns in assemblage attributes changed over time in response to environmental variability, with a tendency for a disruption of upstream-downstream gradients following dry years. 6. Results supported the view that present-day droughts cause relatively small and transient changes to Mediterranean stream fish assemblages. However, longer and more severe droughts expected under altered future climates, may result in declines or local extinctions of the most sensitive species and their potential replacement by more resistant species. Changing drought regimes thus need to be duly considered in the development of conservation strategies for Mediterranean stream fish.
SUMMARY 1. Fish in Mediterranean streams survive through the summer in residual surface waters, encompassing a broad range of abiotic and biotic conditions. Yet, the extent to which fish assemblages may be shaped by functional heterogeneity in dry‐season refugia is largely unknown. This study addresses this issue, by examining fish assemblage and population attributes, and predation patterns in residual summer habitats (12 pools and six runs) across a Mediterranean catchment in south‐west Portugal. 2. Species richness was fairly constant among runs but increased with pool size, with the addition of exotics and rare natives to large pools resulting in nested subsets. The four most common species (chub, nase, loach and eel) were considered generalists in terms of their use of dry‐season refugia. Conversely, rare species presented more specialised habitats, with barbel and exotics favouring pools and stickleback favouring runs. 3. Age and size of the two most common species varied among dry‐season habitats. Age 0 chub were restricted to runs, where spawning stages (age 2 and older fish) were also more represented. Age 0 nase also concentrated in runs, but the individuals collected in pools exhibited greater growth. Conversely, age 2 and older nase were proportionally more abundant in pools, but with greater growth and better condition in runs. 4. The otter was the main fish predator, consuming fish of all species and size classes, irrespective of habitat. Otter activity concentrated in pools, where predation risk for cyprinids seemed to be much higher than in runs. 5. Dry‐season refugia apparently vary in functional importance for different fish species and life stages, acting as complementary units in the landscape. Therefore, the presence of both pool and run refugia trough the summer dry season may play a critical role in promoting the persistence of native species in Mediterranean streams.
Chub Squalius torgalensis and nase Chondrostoma lusitanicum, in a Mediterranean stream, showed important differences in life‐history traits and population dynamics. Both species reached mean maturity at age 2 years. Chub lived up to age 5 years, spawned in March to June, grew at a maximum rate of 0·59 mm mm−1 year−1 and showed a low reproductive allocation, with fecundity and egg size increasing with body size. Nase lived up to age 4 years, spawned in January to April, grew at a maximum rate of 0·46 mm mm−1 year−1 and showed a high reproductive allocation, with egg size independent of body size. Both chub and nase showed moderate fluctuations in population size during 1991–1998, but differed in factors driving density at age. Density of age 1 year juvenile chub decreased following severe summer droughts and proportionate survival prevailed thereafter. Density of age 2 year adult nase decreased following severe spring floods, but neither environmental nor parental stock effects were detected for juveniles and older fishes. The results illustrated the interplay between life history and environmental variability in driving fish population dynamics, with impacts of both summer droughts and spring floods being contingent on species‐specific patterns of spawning and reproductive investment.
Ecological attributes behind the success of the South American cichlid Australoheros facetus in mediterranean-type rivers remain unclear. This study addresses this issue, by analysing its abundance and feeding patterns in the lower Guadiana drainage (Portugal), during the dry season of 2003. Despite slight spatial variations in abundance and individual size, A. facetus seemed well established in the Guadiana, Vascão and Ardila rivers. No seasonal or size-related changes in feeding intensity were found, but the diet changed across rivers, over time and throughout ontogeny. Feeding seemed highly generalist, with fish apparently reacting to both spatial and temporal changes in food supply and tending to use the most abundant and easily accessible food items. Overall prey breadth was broad relative to that of other native and non-native species. It is suggested that a generalist feeding strategy may play a significant role in the establishment of non-native fish in mediterranean-type rivers.
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