Ramon Moreno-Amich

Gradients of life-history variation: an intercontinental comparison of fishes

Multivariate analysis identified atwo-dimensional continuum of life-historyvariation among 301 fish species from Europe,North America, South America and the Atlanticand Pacific coasts of North America. The firstaxis was associated with larger body size,higher fecundity, delayed maturation, fewerreproductive events, and shorter breedingseason on one end and small size, lowfecundity, early maturity, multiplereproductive events per year, and prolongedbreeding season on the other. The second axiscontrasted fishes having larger eggs and moreparental care against fishes with the oppositesuite of traits.Phylogenetic affiliations of species wereapparent in the general patterns of ordinationof species within orders, indicatingevolutionary divergences in life-historypatterns. In fact, partitioning the variance oflife-history traits showed that taxonomic orderand latitude were the most important factorsand geographic region and habitat the least.Despite phylogenetic constraints, basiclife-history patterns showed consistencybetween distantly geographical regions,latitudinal ranges and basic adult habitats,indicating convergences in life-historypatterns. Although the basic life-historypatterns seemed repeatable among distantlyrelated taxa, geographical and latitudinalaffiliations were apparent. Species from SouthAmerica are skewed toward the opportunisticendpoint, whereas North American marine speciesare skewed toward the periodic endpoint of thetrilateral continuum model. Most of the fishspecies from South American data set came fromfluctuating environments, so an opportunisticstrategy of early maturation and continuousspawning permits efficient recolonization ofhabitats over small spatial scales. Incontrast, most species in the North Americanand European data sets came from seasonalhabitats that are nonetheless more hydrologicalstable, so a periodic strategy of delayingmaturation to attain large clutches enhancesadult survivorship during suboptimalenvironmental conditions and recruitment whenearly life stages encounter suitableenvironmental conditions. Similarly,latitudinal affiliations were also observed:opportunistic strategists more common intropical latitudes and periodic strategistsmore common in temperate and Arctic latitudes.

Succession of the animal community in a Mediterranean temporary pond

Abstract Successional changes in the animal community of a temporary pond on the Iberian Peninsula were analyzed based on taxon abundance and the biomass–size spectrum. Changes in the community were studied over 6 hydroperiods during 1996 and 1997. Community dynamics were analyzed using multivariate analysis and a new statistical approach based on the Pareto probability distribution. Three successional phases could be distinguished on the basis of changes in taxon abundance. This pattern is consistent with the 3-phase model reported in other temporary ponds in temperate latitudes. The 1st and 3rd phases corresponded to changes in hydrological factors (pond drying, water turnover) and to seasonality, whereas the 2nd phase corresponded to more stable hydrological conditions. Five successional phases could be distinguished on the basis of biomass–size spectra with similar structure. These phases were consistent with variations over time of parameter c of Pareto Model I. The 3 middle phases (II, III, and IV) identified using the size-spectrum approach corresponded to the single middle phase identified using the taxonomic approach. Thus, community changes during the middle phase were better resolved by analysis of the biomass–size spectrum than by analysis of taxon abundance.

Zooplankton structure and dynamics in permanent and temporary Mediterranean salt marshes : taxon- based and size-based approaches

Differences between zooplankton structure (species abundance and size distribution) and dynamics of permanent and temporary basins of a Mediterranean salt marsh (Emporda Wetlands, NE Spain) were analysed by means of taxon-based and size-based approaches. These basins are shallow bodies of water which are occasionally connected. They were isolated after water retreat from a broadly flooded, low-lying area close to the open sea. Although temporary and permanent basins show no differences in the most abundant zooplankton species, they differ in their zooplankton diversity, temporal pattern and size structure. The zooplankton assemblages of temporary basins exhibit a temporal pattern with six phases, which are conditioned to the hydrological cycle, each one dominated by one species, whereas in the permanent basin this temporal pattern shows only two phases. Zooplankton size distribution of the temporary basins is dominated by large sizes (copepodites and copepods) while the permanent basin is dominated mainly by small sizes (rotifera and nauplii of copepods). In the temporary basins, the shape of the biomass size spectrum changes according to the hydrological cycle and reflects the dynamics of the ecological interactions among zooplankton species. Differences in the zooplankton composition and dynamics of the permanent basin are not a consequence of water permanence but of the ecological interactions among species involved. The presence of a stable fish population in the permanent waters may explain the high values of zooplankton diversity and the low densities of large zooplankton.

Fish zonation in a Mediterranean stream: Effects of human disturbances

Abstract. The fish assemblage and limnological features along twelve sites from the Terri River basin (Catalonia, Spain) were sampled quarterly from August 1999 to May 2000. Twelve fish species were captured, of which four were native and eight exotic. Correspondence analysis revealed that spatial variation accounted for most of the variation (73.7%) in fish species composition. The upstream, urban area was dominated by chub (Leuciscus cephalus) and common carp (Cyprinus carpio) with some exotic species coming from nearby Lake Banyoles, whereas Mediterranean barbel (Barbus meridionalis) predominated in the rest of the Terri River. Along the longitudinal gradient, fish assemblage displayed discontinuous variation and disappearance in more polluted areas. Fish abundance was significantly correlated with summer oxygen concentration and decreased in downstream sampling sites. Species richness was higher in downstream sampling sites due to the proximity of the Ter River. Common carp size increased progressively along the course of the Terri River, whereas the pattern for the eel was the opposite, with the smallest found in downstream sampling sites and a gradual increase in size in the upper reaches. The natural zonation in the fish assemblage of the Terri River is presently altered to a high degree by habitat degradations, pollution, and dispersal of exotic species.

Life-history patterns of 25 species from European freshwater fish communities

Fish life-history patterns were evaluated in relation to the trilateral continuum model by analyzing data from 25 species inhabiting European freshwaters. Multivariate tests identified a trend between later-maturing fishes with higher fecundity, larger size, and few spawning bouts per year and the opposite suite of traits with small fishes. A second trend contrasted fishes having parental care, smaller eggs, and longer breeding seasons against fishes with the opposite suite of traits. As a result, two extreme life-history patterns could be identified among European freshwater fish species: opportunistic and periodic. Nevertheless, intermediate patterns were also present. A true equilibrium life-history pattern was not represented among 25 fish species from European freshwaters. The high concordance of basic life-history patterns among distantly related taxa is probably caused by some universal trade-offs among life-history variables. As a consequence, only a limited life-history patterns may be recognizable among fish species, independently of the origin of fish communities.

Comparison of nutrient and contaminant fluxes in two areas with different hydrological regimes (Empordà Wetlands, NE Spain)

Nutrient (N and P), heavy metal (Ni, Cd, Cr, Cu and Pb) and pesticide (DDT, DDD, DDE, lindane, aldrin, endrin, dieldrin, permethrin, atrazine and simazine) concentrations in water and sediment were analysed in the Empordà Wetlands, a Mediterranean wetland area in NE Spain. Mean nutrient and contaminant concentrations and input and output loads via tributaries were compared in two marshes with different water turnover: a freshwater marsh (FWM), with a high water turnover rate due to continuous surface water inputs and outputs, and a brackish water marsh (BWM), with lower turnover and no continuous surface output, where water remains confined during dry periods. Mean concentrations of most heavy metals exceeded the maximum permissible concentration (MPC) in BWM, whilst only some pesticides reached MPC in FWM. The confined waters of BWM showed higher sensitivity to contaminant input loadings than FWM due to the lack of continuous water outputs. Non-point source pollution inputs during runoff (mainly in FWM) and concentration during confinement (mainly in BWM) showed as the main environmental problems related to nutrients and contaminants in these ecosystems. Thus, the importance of confinement and its effect on pollutant concentrations must be borne in mind to achieve correct management of Mediterranean wetlands.

Quantifying the activity and movement of perch in a temperate lake by integrating acoustic telemetry and a geographic information system

The horizontal movement patterns of perch (Perca fluviatilis L.) were studied by means of an automatic positioning system in a small Mediterranean temperate lake in northeastern Spain. Seven perch (247–265 mm in fork length, 243–271 g in weight) were implanted surgically with ultrasonic transmitters (38 mm long, 3.5 g in water) and their positions monitored approximately every 180 s using a radio-linked acoustic positioning (VRAP) system. Fish positions were integrated into a geographic information system database with information on date, time, slope and bottom depth. As a measure of the intensity of movement, the swimming speed was calculated. The Minimum Convex Polygon area was obtained as a measure of the area of activity. The data dispersion was calculated as r 2 (the mean squared distance from the center of activity). The tagged perch showed individual variations in activity levels, but all had a circadian pattern, being less active at night (average speed 6.0 cm s−1) than during the day (average speed 10.4 cm s−1). They exhibited two peaks of activity, at sunrise and sunset. Two patterns of movement emerged, the first characterised by movement between the limnetic and the littoral zones, the second restricted to the shoreline. Fish appeared to follow the same path, reducing the area of activity to a limited part of the lake. Also, fish returned to the same place after movements, showing site fidelity but only for a few days. Overall, although perch could be very abundant in open waters (limnetic zone), they demonstrated a high dependence on the littoral zone.

Rudd (Scardinius erythrophthalmus) introduced to the Iberian peninsula: feeding ecology in Lake Banyoles

The first data on the ecology of rudd (Scardinius erythrophthalmus) introduced to the Iberian peninsula are presented. The habitat and diet variation of rudd were studied in Lake Banyoles (Spain), an oligotrophic karstic lake dominated by exotic fish species. Rudd were strictly littoral and the diet was based on detritus and plant material. The most important animal prey were the cladocerans Daphnia longispinaand Scapholeberis rammneri, amphipods and several late stages of nematoceran dipterans. Rudd were more zooplanktivorous in spring and autumn and less in summer. There was also a size-dependent diet shift, from microcrustaceans to macroinvertebrates. The diet of rudd was also distinguished by the importance of plant material and various small neustonic invertebrates, particularly S. rammneriand late stages of nematocerans, showing a strong resource partitioning with other fish species. The degree of herbivory in Lake Banyoles was lower than usual.

Population dynamics of Triops cancriformis (Crustacea: Branchiopoda: Notostraca) of the Espolla temporary pond in the northeastern Iberian peninsula

The population dynamics of Triops cancriformis in Espolla temporary pond (NE Iberian peninsula) were studied during 1996 and 1997, which encompassed six flooded periods. Data were collected on each individuals size, sex, and, if female, on number of eggs in the oostegopodes. Male-biased sex ratios were found only in the drying-out phase and variations in fecundity were strongly related to hydroperiod duration. Sex ratio variation during the drying-out phase can be attributed to female mortality because the very low recruitment observed does not support the hypothesis of an increase of males. Two hypotheses are advanced to account for female mortality: (1) differential reproductive effort, and (2) size selective predation by herons. This population is characterised by low values of maximum densities compared with other notostracan populations, and by higher densities in the spring–summer hydroperiods than in the winter ones.

Seasonal abundance and depth distribution of Blennius fluviatilis and introduced Lepomis gibbosus, in Lake Banyoles (Catalonia, Spain)

Temporal and spatial occurrence of limnetic larval fish in Lake Banyoles in the spring and summer of 1993 and 1994 indicated that pumpkinseed spawned over 15 weeks (from mid-May to mid-August) and freshwater blenny over 14 weeks (from mid-May until early August). We noted that the spawning period of pumpkinseed commenced earlier in Lake Banyoles than in more northern lakes.Pumpkinseed and freshwater blenny larvae were collected by tow net in both the limnetic (at 4–4.5 m) and littoral zones (at 0–0.5 m), during daylight (no samples were taken during night). Freshwater blenny larvae were more abundant than pumpkinseed larvae. Both of them were more abundant in 1993 than in 1994 (pumpkinseed: 1.61 ind. 100 m−3 vs. 0.70 ind. 100 m−3; freshwater blenny: 3.95 ind. 100 m−3 vs. 1.90 ind. 100 m−3) and in both years they were uniformly distributed in the lake, without differences between the areas considered. The pumpkinseed larvae migrated at 3.5 mm TL into the limnetic zone, and returned to the vegetation of the littoral zone as juveniles. The freshwater blenny larvae were also planktonic (3.5–14 mm TL) but were distributed in both the littoral and limnetic zone before benthic settlement. The larval distribution is discussed on the basis of a trade-off between foraging and predation risk.