Luís Cancela da Fonseca

Analysis of competitive interactions between the limpets Patella depressa Pennant and Patella vulgata L. on the northern coast of Portugal

Inter- and intraspecific competitive interactions among the species Patella depressa Pennant and P vulgata L. were analysed on the northern coast of Portugal, where both species co-occur in similar proportions. Increased (x 2, x 4), normal and decreased (x 1/2 intraspecific only) densities of limpets were used to test the effects of competition on the growth and mortality of the limpets, and competitive interactions between the different species. Fenced plots of 25 x 25 cm enclosing marked limpets at various densities were set up at a mid tidal level on the shore. Twelve treatments with three replicates of different combinations of densities and species were established. Mortality was recorded every 15 days and length was measured monthly during the 5 months of the experiment. At the end of the experiment, limpets were collected for biometry, sex determination and gonad staging. Both species of grazing molluscs showed increased mortality and reduced size and weight in increased density treatments. Limpets in decreased density treatments showed lower mortality and higher size. There were no significant differences between the effect of P. vulgata on R depressa and the effect of P. depressa on P. vulgata on mortality and length. There were, however, decreases in weight due to intraspecific effects for both species. For P. vulgata, interspecific effects were much less than intraspecific effects and for P. depressa, the interspecific effects depended on density. The analysis of the gonad stage at the end of the experiment showed that the lower stages of development were most prevalent at increased densities and the occurrence of neuter individuals only in increased densities single species plots corroborated the evidence of a stronger intraspecific interaction. The availability of food was indirectly assessed by determination of chlorophyll concentration with spectrophotometric analysis of rock chips. Microalgal food assessment in this experiment, however, did not show any clear trends. The results were compared with previous studies of competition in grazing molluscs and discussed in relation to the limits of distribution of P vulgata on the Portuguese coast.

The effects of grazing on the distribution and composition of low-shore algal communities on the central coast of Portugal and on the southern coast of Britain

A red algal turf is often found just below the barnacle/limpet zone of many European shores, especially on steep shores of moderate exposure. The hypothesis that grazing by limpets determines the upper limit of distribution of this red algal turf was tested on moderately exposed shores in Portugal and Britain. We also aimed to assess whether the grazing effect is modified at various spatial scales. Grazers were excluded by fences, with half-fenced and unfenced controls. Exclusion plots were rapidly colonised by green ephemeral algae in the months immediately after the beginning of the experiment (summer); these algae were later replaced by perennial algae. The percentage cover of turf-forming macroalgae showed a significant increase at both locations. The upper limit of distribution extended more than 50 cm on most of the shores studied. In contrast, control and half-fenced plots remained devoid of algae. After 2 years, ungrazed plots were mainly colonised by a red algal turf (e.g. Caulacanthus ustulatus, Gelidium spp., Laurencia pinnatifida) in Portugal, while canopy cover (Fucus serratus and Himanthalia elongata) dominated in Britain in marked contrast to the grazed plots. Physical factors acting at both local and geographical scales may explain these differences. However, although physical factors probably have an important influence on the identity, size and abundance of sublittoral fringe macroalgae, grazers play a major role in directly setting their upper limits. The effect of grazing by limpets was not consistent for all of the morphological algal groups and spatial scales considered in the present study. The effect of grazing on the cover of turf algae varied between Portugal and Britain (location scale), while effects on ephemeral and canopy algal cover varied at the shore scale within location.

The relationship between phytoplankton diversity and community function in a coastal lagoon

The decrease of biodiversity related to the phenomena of global climate change is stimulating the scientific community towards a better understanding of the relationships between biodiversity and ecosystem functioning. In ecosystems where marked biodiversity changes occur at seasonal time scales, it is easier to relate them with ecosystem functioning. The objective of this work is to analyse the relationship between phytoplankton diversity and primary production in St. Andre coastal lagoon — SW Portugal. This lagoon is artificially opened to the sea every year in early spring, exhibiting a shift from a marine dominated to a low salinity ecosystem in winter. Data on salinity, temperature, nutrients, phytoplankton species composition, chlorophyll a (Chl a) concentration and primary production were analysed over a year. Modelling studies based on production-irradiance curves were also conducted. A total of 19 taxa were identified among diatoms, dinoflagellates and euglenophyceans, the less abundant group. Lowest diversities (Shannon— Wiener index) were observed just before the opening to the sea. Results show a negative correlation (p 90% of cell abundance) and production was maximal (up to 234.8 mg C m−3 h−1). Maximal photosynthetic rates (Pmax) (2.0–22.5 mg C mg Chl a −1 h−1) were higher under lower Chl a concentrations. The results of this work suggest that decreases in diversity are associated with increases in biomass and production, whereas increases correspond to opposite trends. It is suggested that these trends, contrary to those observed in terrestrial and in some benthic ecosystems, may be a result of low habitat diversity in the water column and resulting competitive pressure. The occurrence of the highest photosynthetic rates when Chl a is low, under some of the highest diversities, suggests a more efficient use of irradiance under low biomass-high diversity conditions. Results suggest that this increased efficiency is not explained by potential reductions in nutrient limitation and intraspecific competition under lower biomasses and may be a result of niche complementarity.

Phylogeny, molecular ecology and taxonomy of southern Iberian lineages of Triops mauritanicus (Crustacea: Notostraca)

We investigated the phylogeography of the main lineages in the tadpole shrimp Triops mauritanicus Ghigi in the south-western Iberian Peninsula, using mitochondrial 12S and 16S rDNA sequences. Our results indicate that a fourth, hitherto unknown main phylogenetic lineage occurs in Iberia, so that in total, the species is divided into six distinct clades, comprising T. m. mauritanicus, T. m. simplex Ghigi, and four as yet unnamed lineages that appear to be endemic to Iberia. Percentages of sequence divergence among the main clades in T. mauritanicus reach the range reported for recognized species in other notostracan lineages. A thorough morphological investigation also revealed that the differentiation among these lineages is higher than previously thought, and that populations of three of the main clades within T. mauritanicus can be reliably separated from each other and from the remaining lineages based on the morphology of adult males. The remaining clades also show a significant level of morphological differentiation, but include a certain proportion of populations for which the additional application of molecular methods is needed for a reliable determination. The geographic distributions of 12S haplotypes are indicative of frequent dispersal events and gene flow among populations belonging to the same main lineage, but give no evidence of recent migration events among different main lineages, suggesting that there is no gene flow among the latter. Our data thus suggest that the six main lineages within T. mauritanicus represent distinct species. We therefore describe the Iberian lineages as T. baeticus Korn n. sp., T. emeritensis Korn & Pérez-Bote n. sp., T. gadensis Korn & García-de-Lomas n. sp., and T. vicentinus Korn, Machado, Cristo & Cancela da Fonseca n. sp., and reinstate T. simplex Ghigi to full species status. Our data confirm the general, previously recognized pattern of a lower dispersal probability in gonochoric Triops taxa. However, we found evidence that passive dispersal in Triops may be further complicated by a strong habitat dependence of dispersal probability, mediated by prevailing dispersal vectors.

Sister species within the Triops cancriformis lineage (Crustacea, Notostraca)

We investigated the phylogenetic relationships among the three presently recognized subspecies of the tadpole shrimp, Triops cancriformis, using mitochondrial 16S and 12S rDNA sequences. Our results indicate that the taxon is divided into two distinct lineages. One lineage is formed of T. c. cancriformis populations and samples from northern Spain that had been classified as T. c. simplex in the most recent literature. The second lineage comprises all populations of T. c. mauritanicus and northern African populations of T. c. simplex. These two main lineages separated 2.3 to 8.9 million years ago, based on the range of inferred molecular clocks recognized for crustacean mtDNA sequence divergence. Percentages of divergence are in the range reported for recognized species in other notostracan lineages and we therefore propose to recognize them as two species, Triops cancriformis and Triops mauritanicus. The latter would comprise two subspecies in northern Africa, one consisting of the Moroccan populations of the former T. c. mauritanicus, the other comprising the African populations of the former T. c. simplex. It also includes three as‐yet unnamed lineages. A comparison of morphological characters with the molecular data revealed that the former T. c. simplex cannot be reliably separated from T. c. cancriformis, using morphological characters that have hitherto been used to distinguish among subspecies of T. cancriformis. Our investigation is the first to demonstrate the presence of T. c. cancriformis in Africa (Tunisia). The genetic haplotypes of these populations are identical with haplotypes also occurring in Central and Western Europe, as well as in Sicily. Therefore, we hypothesize that the African populations of T. c. cancriformis represent a result of repeated long‐distance dispersal across the Mediterranean Sea.

Trophic structure of macrobenthic communities on the Portuguese coast. A review of lagoonal, estuarine and rocky littoral habitats

Abstract This work is based on a compilation and treatment of data obtained on several studies regarding the macrobenthos trophic structure, carried out in different habitats of the Portuguese coast: a lagoon, rocky subtidal shores and estuarine eelgrass beds. Macrobenthic organisms were assigned to four distinct trophic groups (filter feeders, detritivores, herbivores and carnivores). Detritivores were the dominant trophic group in soft-bottom communities. Filter feeders dominated in subtidal rocky shores while eelgrass communities were equally represented by detritivores and herbivores. Current intensity and sediment deposition are discussed as factors affecting the observed distribution. Nevertheless, different sampling methods used in these studies and the lack of information on feeding habits of some species can also influence the results obtained.

Macrobenthic Colonisation of Artificial Reefs on the Southern Coast of Portugal (Ancão, Algarve)

Artificial reef systems play an important role in the increase of natural production of biological marine resources and they have been deployed worldwide. In Portugal, seven artificial systems have been deployed along the southern coast of the Algarve. Research to date has focussed mainly on fish populations, particularly those of economical importance. The present work aims to study the macrobenthic communities of the artificial reef structures, as these will enhance the food resources and shelter, making the reefs more attractive to fish. In particular, we experimentally analysed the sequence of colonisation of macrobenthic communities of the Ancao artificial reef system, which was deployed in the summer of 2002. The study of the colonisation of benthic communities was done using samples of concrete cubic units (15 × 15 cm) that were suspended at the reef modules at a depth of 20 m, at the time of the reef immersion. Four replicate samples were collected by SCUBA diving from two groups of the Ancao reef every three months from the starting date. Sampling was done using essentially non-destructive methods to assess the percentage cover of macrobenthic organisms in both vertical and horizontal surfaces. The percentage cover of the taxonomic groups was compared within the different surfaces of the samples and between the two reef groups. The bottom surface of cubic samples had a significantly higher colonisation related to the dominance of barnacle cover, probably due to lower sedimentation levels. Samples from both reef groups showed a similar pattern of colonisation. Barnacles, bryozoans and serpulids dominated the samples three months immediately after the beginning of the experiment. Other invertebrates groups, such as Porifera, Hydrozoa, Anthozoa, other sessile Polychaeta, Decapoda, Gastropoda and Bivalvia, were more abundant after six months of colonisation.

How complementary are epibenthic assemblages in artificial and nearby natural rocky reefs

The present study analyses the composition, structure and trophic function of epibenthic assemblages in two artificial reefs (ARs) 16 years after deployment and in nearby natural reefs (NRs), aiming at providing insights on the complementarity between both habitats. Current findings suggest that after 16 years the ARs (concrete blocks), located in southern Portugal, do not act as surrogates for NRs, as epibenthic assemblages differed between reef types in composition, structure and trophic function. NRs showed higher diversity and complementarity (i.e. beta-diversity) than ARs, evidencing higher redundancy. Higher heterogeneity within NRs was also evidenced by the multi-dimensional scaling analysis based on abundance, biomass and trophic composition. NRs presented higher abundance of molluscs and biomass of sponges, resulting in differences in the trophic function: suspension-feeding dominated the NRs, while within ARs there was an ascendency of carnivory. Although not acting as surrogates for NRs and provided that no adverse effects (e.g. establishment of non-native species) were detected, ARs may have a significant contribution for the increase of regional diversity, as evidenced by the highest complementarity levels observed between assemblages in both reefs.

Ecological quality assessment of small estuaries from the Portuguese coast based on benthic macroinvertebrate assemblages indices.

Benthic macroinvertebrates communities are the most consistently emphasized biotic component of aquatic ecosystems and are one of the biological indicators required for assessment by the European Water Framework Directive. In this context, several indices based on these communities have been developed in order to assess ecological quality of estuarine systems. In the present work we used AMBI, M-AMBI, BENTIX and BAT to distinguish ecological status of five small estuarine systems of the Portuguese south and southwest coasts. Although indices outputs did not differ between systems and sampling seasons, results indicated that the metrics in which these indices are based could differentiate community structures as a result of two main gradients that force these communities: the natural variability, and the anthropogenic impact.

A coupled biogeochemical-Dynamic Energy Budget model as a tool for managing fish production ponds

The sustainability of semi-intensive aquaculture relies on management practices that simultaneously improve production efficiency and minimize the environmental impacts of this activity. The purpose of the present work was to develop a mathematical model that reproduced the dynamics of a semi-intensive fish earth pond, to simulate different management scenarios for optimizing fish production. The modeling approach consisted of coupling a biogeochemical model that simulated the dynamics of the elements that are more likely to affect fish production and cause undesirable environmental impacts (nitrogen, phosphorus and oxygen) to a fish growth model based on the Dynamic Energy Budget approach. The biogeochemical sub-model successfully simulated most water column and sediment variables. A good model fit was also found between predicted and observed white seabream (Diplodus sargus) growth data over a production cycle. In order to optimize fish production, different management scenarios were analysed with the model (e.g. increase stocking densities, decrease/increase water exchange rates, decrease/increase feeding rates, decrease phosphorus content in fish feeds, increase food assimilation efficiency and decrease pellets sinking velocity) to test their effects on the pond environment as well as on fish yields and effluent nutrient discharges. Scenarios were quantitatively evaluated and compared using the Analytical Hierarchical Process (AHP) methodology. The best management options that allow the maximization of fish production while maintaining a good pond environment and minimum impacts on the adjacent coastal system were to double standard stocking densities and to improve food assimilation efficiency.