Age determination in Pomatoschistus mi nut us (Pallas) and Pomatoschistus microps (Kreyer) (Pisces: Gobiidae) from the upper Tagus estuary, Portugal

The use of scale inspection as a means of age-determination in Pomatoschistus microps and Pomatoschistus minutus in the Tagus estuary is confirmed, but annulus formation pattern is different from that described for P. microps in the area of the British Isles (Miller, 1975). In Britain, two annuli are laid down on the scales each year, but in the Tagus, only one annulus is formed. One possible reason for this difference is the fact that in the Tagus the peak breeding season and the lowest water temperatures coincide. Therefore, the two causes of annulus formation coexist at the same time of year.
Population structure in both species is dominated by the 0-group, which appears earlier (April/May) than reported for other areas. Maximum age seems not to exceed 26 months in P. microps and 32 months in P. minutus .     

Composition, temporal changes and ecological guild classification of the ichthyofaunas of large European estuaries – a comparison between the Tagus (Portugal) and the Elbe (Germany)

Fish species densities from 1025 hauls obtained with beam trawl and stow net catches from the southern Tagus and northern Elbe estuaries were analysed for two different periods of time between 1978 and 1997. The data were related with environmental factors to compare species composition and densities, contributions of different ecological guilds, structuring environmental key factors, intra‐annual cyclicity and interperiodical stability of ichthyofaunas in large European estuaries at different latitudes. Although the total number of species in both the Tagus and Elbe estuaries (Tagus: 57 species, Elbe: 58 species) was very similar, significant differences were estimated in species compositions, species densities and number of species, and individuals of the different ecological guilds between both estuaries. A total of 15 species occurred in both estuaries but only two of these species, the catadromous Anguilla anguilla and the marine estuarine opportunist Pleuronectes flesus, contributed substantially to the total abundance. Marine species together contributed more than 5% of the total number of species in the Tagus and Elbe estuaries, whereas their contribution in number of individuals amounted to 22% in the Tagus and only 9% in the Elbe. In terms of number of individuals, the ichthyofauna of the Tagus estuary was dominated by estuarine species (especially Pomatoschistus spp.) contributing 68% of the total abundance. In contrast, the fish fauna of the Elbe estuary was dominated by high numbers of anadromous species (88%) individuals, especially Osmerus eperlanus. Canonical corresponding analyses (CCA) indicated that, besides intra‐annual variables, salinity and mouth distance were two of the most important environmental factors structuring the fish communities in both estuaries. Moreover, water depth was found to be an important environmental factor in the Tagus estuary, whereas the water temperature substantially affected the ichthyofaunal composition in the Elbe estuary. In both estuaries, contributions by number of species and number of individuals of the different ecological guilds were not statistically different between the two periods considered, 1978–80 and 1995–97 for the Tagus, and 1981–86 and 1989–95 for the Elbe. Nevertheless, from 1981–86 until 1989–95 a remarkable increase of about 4.5 times in the densities of O. eperlanus was observed in the Elbe estuary, probably related to improved water quality, especially of oxygen conditions in the nurseries since 1990. This was also supported by the results of CCA, suggesting important influences of the years 1989 and 1990 on the Elbe estuary fish assemblage. Climate change as a result of global warming may be evident for the observed changes of the densities of P. flesus which decreased in the southern Tagus estuary and, in contrast, increased in the northern Elbe estuary during comparable periods of time. It is recommended to use the ecological guild classification scheme which was applied for the Tagus and Elbe estuaries for further studies of European estuarine fish assemblages.     

Spatial Patterns of Benthic Macroinvertebrates in Intertidal Areas of a Southern European Estuary: The Tagus, Portugal

This study characterizes the composition and spatial distribution patterns of the benthic macrofauna in the intertidal mudflats of the Tagus estuary, western Portugal. A total of 68 species, more than 226,000 specimens with a total wet weight biomass of approximately 1170 g were identified in 380 sites. The species Streblospio shrubsolii, Cyathura carinata, Tharyx sp., Hydrobia ulvae and Tubificids were the most common and abundant. Scrobicularia plana strongly dominated the biomass. The invertebrate macrofauna of the Tagus estuary shows similarities to what is known from other temperate mudflats. The diversity of species, their overall abundance and the ratio of Molluscs plus Crustaceans to Polychaete species corroborate the distinctiveness between temperate and tropical mudflats and sandflats. The spatial distribution of the fauna reflects the sediment characteristics but the relationship between the environmental and the biological data is not as strong as obtained for sublittoral areas. This relationship diminishes from the sublittoral shelf to sublittoral estuarine areas, showing minimum values in this study, suggesting that such a relationship is less straightforward as natural disturbance increases. Nevertheless, a mixture of grain-size, elevation (inundation time) and particular habitats (relic oyster beds) form the best explanatory factors for the spatial distribution patterns of the intertidal benthic macrofauna of the Tagus estuary.     

Four new species of Gyrodactylus von Nordmann, 1832 (Monogenea,Gyrodactylidae) on gobiid fishes: combined DNA and morphological analyses

Four Gyrodactylus species parasitising four closely related gobiid species in European coastal waters were studied and compared with G. arcuatus Bychowsky sensu Bychowsky & Poljansky (1953) from Gasterosteus aculatus . These were G. gondae n. sp. from Pomatoschistus minutus and P. lozanoi , G. flavescensis n. sp. from Gobiusculus flavescens , G. arcuatoides n. sp. from P. minutus and G. branchialis n. sp. from P. microps. Combined molecular and morphological analyses, as well as morphometric and statistical methods, were used. The ssrRNA V4 region and the complete ITS rDNA region were sequenced. Genetically the four new species are clearly distinct from G. arcuatus . From a morphological point of view, the haptoral hard parts of G. gondae n. sp., G. flavescensis n. sp. and G. arcuatoides n. sp. are related to those of G. arcuatus, while these parts of G. branchialis n. sp. are different, but related to those of G. quadratidigitus Longshaw, Pursglove & Shinn, 2003. For the latter two species, a new species group is formed. The V4 and ITS sequence analyses, however, indicate a close relationship between G. branchialis and the three G. arcuatus-like species.     

Organotin in the Tagus estuary

The tri(n-butyl)tin (TBT) moiety has been shown to have complex ecotoxicological effects on estuarine populations. In the Tagus the massive die-off of the Portuguese oyster (Crassostrea angulata Lmk), has been attributed to the introduction and use of this contaminant by shipyards. This paper reviews previous work by others and reassesses our results obtained in water and sediments of the Tagus estuary, as a contribution to the understanding of tin geochemistry and organotin speciation in this estuary. Contrary to earlier results, the latest surveys show that in the open waters of the Tagus measurable concentrations of all the three butyltin species occur, exhibiting peak concentrations at two locations that indicate two previously undetected sources of TBT. TBT also presents a peak concentration in sediments near the Lisnave shipyards, as previously suggested. The latest surveys still detected methylated forms of tin, particularly monomethyltin (Me Sn³⁺), in Tagus sediments, with maximum concentrations in the vicinity of urban effluent discharges. The concentrations for TBT and its degradation products found in the Tagus are potentially harmful to the populations of gastropods and endemic bivalves.     

Importance of discards of a beam trawl fishery as input of organic matter into nursery areas within the Tagus estuary

The brown-shrimp beam trawl fishery carried out within the Tagus estuary produces discards due to the little commercial interest of most of the species caught. Between 1994 and 1996, monthly samples were collected in the two major fishing areas within the Tagus estuary, using a beam trawl, in order to estimate the amount of fish and crustaceans caught per unit of effort. The fishing effort of the commercial fleet was determined based on surveys of professional fishermen. Mortality estimates of discards were also evaluated experimentally. The main fish and crustacean species discarded after capture were Crangon crangon (Linnaeus, 1758), Liza ramada (Risso, 1826), Carcinus maenas (Linnaues, 1758) and Pomatoschistus minutus (Pallas, 1770). The estimated total amount of fishery discards in the upper part of the Tagus estuary is approximately 1500 tonnes per year, which represents ca. 90% of the captures. The mortality rate of the fishes and crustaceans discarded varied according to species and season, with the highest rates during Summer months. Considering nitrogen and carbon content of the main discarded species, an input of particulate organic matter of more than 140 tonnes of carbon and 35 tonnes of nitrogen per year were estimated for these estuarine areas.     

Differential drift and parasitism in invading and native Gammarus spp. (Crustacea: Amphipoda)

Invading and native species often interact directly, such as by predation, producing patterns of exclusion and coexistence. Less direct factors, such as interactions with the broader abiotic and biotic environment, may also contribute to such patterns, but these have received less recognition. In Northern Ireland, the North American Gammarus tigrinus has invaded freshwaters populated with the native Gammarus duebeni celticus , with intraguild predation between the two implicated in their relative success. However, these species also engage in day and night "drifting", an activity that subjects amphipods to intense predation from fish and wildfowl. Sampling of two rivers where the invader and native co-occur showed that, compared with the benthos, G. tigrinus was underrepresented and G. d. celticus overrepresented in the drift. In addition, G. tigrinus were free from parasites, whereas some G. d. celticus harboured the acanthocephalans Polymorphus minutus and Echinorhynchus truttae and the muscle wasting microsporidian Pleistophora sp. (new species). Compared with the benthos, G. d. celticus parasitized with P. minutus were overrepresented and unparasitized individuals underrepresented in the drift. The opposite pattern was found with Pleistophora sp . In laboratory experiments, G. tigrinus were less positively phototropic and less "active" than G. d. celticus (unparasitized animals). Polymorphus minutus increased G. d. celticus positive phototropism and activity, while Pleistophora sp. increased positive phototropism but decreased activity. Previous studies show that the invader G. tigrinus is more disadvantaged by intraguild predation from the native G. d. celticus than vice versa. However, the native appears more disadvantaged with respect to drift, parasitism and the interaction of the two. These factors may mitigate direct interactions and help explain complex patterns of coexistence between these invader and native species.     

Species zonation in Corroios salt marsh in the Tagus estuary (Portugal) and its dynamics in the past fifty years

The objective of this study was to understand the main factors controlling salt marsh plant species structure and dynamics. So, we determined plant cover and composition across a permanent transect, 450 m long and 1 m wide, defined in 1951 in Corroios salt marsh, in the Tagus estuary (Portugal) and we characterized the physicochemical variables every 50 m along this transect. Based on those results we discuss the dynamic and evolution of salt marsh vegetation during the last 50 years comparing former and recent data. The results showed that differences in salinity and flooding were determinant factors in plant species composition and distribution along the studied transect. In addition, long-term variations of these factors as a consequence of vertical accretion and sea level rise seem to be responsible for the evolution in plant structure and vegetation zonation patterns, during the last 50 years in the Tagus estuary salt marshes.     

Identification of a host-associated species complex using molecular and morphometric analyses,with the description of Gyrodactylus rugiensoides n. sp. (Gyrodactylidae,Monogenea)

Gyrodactylus rugiensis was originally described as a parasite occurring on the marine gobies Pomatoschistus minutus and Pomatoschistus microps. In our preliminary survey this species was also frequently found on Pomatoschistus pictus and Pomatoschistus lozanoi. Subsequent molecular analysis of the internal transcribed spacers rDNA region revealed that this parasite actually represents a complex of two apparently cryptic species, one restricted to P. microps and the other shared by P. minutus, P. lozanoi and P. pictus. Morphometric analyses were conducted on 17 features of the opisthaptoral hard parts of specimens collected from all four host species. Standard discriminant analysis showed a clear separation of both genotypes by significant differences in marginal hook and ventral bar features. Statistical classifiers (linear discriminant analysis and nearest neighbours) resulted in an estimated misclassification rate of 4.7 and 3.1%, respectively. Based on molecular, morphological and statistical analyses a new species, Gyrodactylus rugiensoides is described. This species seems to display a lower host-specificity than generally observed for Gyrodactylus species as it infects three sympatric host species.However, seasonal and host-dependent morphometric variation is shown for G. rugiensoides collected on P. pictus. Host-switching and gene flow might be important factors preventing speciation on closely related and sympatric host species. The presence of host associated species complexes in this Gyrodactylus-Pomatoschistus system is also confirmed by the presence of two host-dependent genotypes within G. micropsi found on P. minutus and P. lozanoi, and P. microps, respectively. By comparing host and parasite phylogeny, phylogenetic and ecological factors influencing host-specificity are discussed.     

Polymorphus minutus affects antitoxic responses of gammarus roeseli exposed to cadmium

The acanthocephalan parasite Polymorphus minutus is a manipulator of its intermediate host Gammarus roeseli, which favours its transmission to the final host, a water bird. In contaminated environments, G. roeseli have to cope with two stresses, i.e. P. minutus infection and pollutants. As P. minutus survival relies on its host's survival, we investigated the influence of P. minutus on the antitoxic defence capacities and the energy reserves of G. roeseli females after cadmium exposure. In parallel, malondialdehyde, a toxic effect biomarker, was measured in G. roeseli females and in P. minutus. The results revealed that infected females displayed higher cell damage than uninfected ones, despite an apparent increase in reduced glutathione and metallothionein production. In fact, the increase of these antitoxic systems could be counterbalanced by carotenoid intake by the parasite, so that the overall defence system seemed less efficient in infected females than in uninfected ones. In addition, we demonstrated that cadmium induced cell damage in P. minutus, probably linked with cadmium accumulation in the parasite. Altogether, we observed a paradoxical pattern of responses suggesting that P. minutus increases cadmium toxicity in G. roeseli females although (i) it tends to increase several host antitoxic defence capacities and (ii) it bears part of the pollutant, as reflected by cell damage in the parasite.     

Spatial and temporal variation of physico-chemical conditions and phytoplankton during a dry year in the Tagus estuary (Portugal)

During monthly synoptic surveys in a dry year, physicochemical and biological data were collected at 15 stations throughout the Tagus estuary. A Principal Component Analysis (PCA) was used to identify the major sources of variation in the data. The most important variations in the Tagus estuary were the spatial variations, mainly the longitudinal ones showing clearly the riverine and the sea water influenced zones of the estuary. The results also showed that a lateral variation was due to the local pollution as indicated by intense organic matter degradation and high rates of oxygen consumption. The seasonal variation related to temperature and the variation associated with the semidiurnal tidal cycle were also apparent. These, however, contributed less than the spatial variation to the total variability in the measured parameters of the Tagus estuary.     

On the use by birds of intertidal areas of the Tagus estuary: implications for management

The use of intertidal areas of the Tagus estuary by birds was re-analysed, based on data from 1990 to 1993, to describe (i) the temporal and spatial patterns of use of intertidal areas, (ii) the diet of birds during winter and (iii) the habitat selection patterns of feeding birds, during winter. The most common birds in the estuary were gulls, waders and ducks. Highest densities were recorded for most species in autumn and winter. The spatial distribution of birds in the intertidal areas of the estuary did not vary significantly across seasons, although broader distributions occurred when bird populations were present in high numbers. In autumn, use of intertidal areas was highly variable. Specific areas in the estuary were identified as holding important densities of birds, or having a high species richness. The most abundant species selected feeding areas according to sediment type, although the presence of channels, saltmarsh or humans also influenced the distribution of birds. The invertebrates Carcinus maenas, Hediste diversicolor and Scrobicularia plana were identified as key prey species. Plant material was important as food for ducks. Implications of these findings for the management of the estuary are discussed.     

Short-term sedimentation in Tagus estuary,Portugal: the influence of salt marsh plants

Short-term sediment deposition was studied at four salt marsh areas in the Tagus estuary. In areas covered with Sarcocornia perennis, Sarcocornia fruticosa, Halimione portulacoides and Spartina maritima and also in the non-vegetated areas, sedimentation was measured as the monthly accumulation of sediments on nylon filters anchored on the soil surface, from August 2000 to May 2001. Our experiments were used also to determine the influence of the different plant species in vertical accretion rates. Short-term sedimentation rates (from 2.8 to 272.3 g m⁻² d⁻¹) did show significant differences when the four salt marshes studied in the Tagus estuary were compared to each others. Salt marshes closer to the sediment sources had higher sedimentation rates. Our results suggest that the salt marsh type and surface cover may provide small-scale variations in sedimentation and also that sediment deposition values do change according to the position of the different plant species within the salt marsh. Sedimentation is an essential factor in salt marsh vertical accretion studies and our investigation may provide support to help forecast the adaptative response of the Tagus estuary wetlands to future sea level rise.     

Paleoclimatic history and vicariant speciation in the "sand goby" group (Gobiidae, Teleostei)

Vicariant and climatic cycling speciation hypotheses of the 'sand gobies' belonging to the genera Pomatoschistus, Gobiusculus, Knipowitschia, and Economidichthys are tested using molecular phylogenies constructed of nuclear DNA (ITS1 locus) and mitochondrial DNA (12S and 16S fragments). These gobies are among the most abundant in the Eastern Atlantic-Mediterranean region, and play an important role in the ecosystem. Considerable ITS1 length differences, primarily due to the presence of several tandem repeats, were found between species and even within individuals. Therefore, phylogenetic analyses focused on fragments of the 12S and 16S mtDNA region that have been sequenced for 16 goby taxa. The 'sand gobies' clustered as a monophyletic group as proposed on morphological grounds. However, G. flavescens, E. pygmaeus, and K. punctatissima clustered within the Pomatoschistus species, pointing to a paraphyletic origin of these genera. Furthermore, the genetic divergence between P. minutus from the Adriatic Sea versus the Atlantic-Mediterranean region was as high as the divergence within the P. minutus complex, suggesting that P. minutus from the Adriatic Sea should be considered as a distinct species. The "star" phylogeny might suggest that these gobies evolved in a very short time period, possibly linked to the drastic alterations in the Mediterranean Sea during and immediately after the Messinian salinity crisis at the end of the Miocene. The freshwater life-style appeared monophyletic; equating its origin with the salinity crisis resulted in a molecular clock estimate of 1.4% divergence per million years. The last common ancestor probably occupied sandy bottoms and a coastal niche while several species subsequently adapted to new habitats (pelagic, freshwater or stenohaline). The origin of the shallowest clades dated back to the glacial cycling during the Pleistocene epoch.     

Southern limit of distribution of the soft-shell clam <Emphasis Type="Italic">Mya arenaria</Emphasis> on the Atlantic East Coast

The invasive soft-shell clam Mya arenaria was recorded for the first time in the Tagus estuary, central Portugal. Propagules of the soft-shell clam were probably unintentionally introduced into the Tagus estuary through ballast water. Mya arenaria was observed in muddy sands in the upper intertidal zone. The assemblage in which the soft-shell clam was observed was composed of typical estuarine species such as the gastropod Hydrobia ulvae, the bivalve Scrobicularia plana and several polychaete species. The population density of Mya arenaria in the finding area was 40 ind m⁻² with a fresh biomass of 163.5 g FW m⁻².     

Carotenoids of two freshwater amphipod species (Gammarus pulex and G. roeseli) and their common acanthocephalan parasite Polymorphus minutus

Carotenoid compositions of two freshwater Gammarus species (Crustacea: Amphipoda) and of their common acanthocephalan parasite Polymorphus minutus were characterized. The effect of carotenoid uptake by the parasite was addressed by comparing the carotenoid content of uninfected and infected female hosts. Using high-pressure liquid chromatography (HPLC), co-chromatography of reference pigments and electron ionization mass spectrometry of collected HPLC fractions (EI-MS), several xanthophylls and non-polar compounds were identified. Seven kinds of carotenoids, mainly xanthophylls, were identified in gammarids. Astaxanthin was predominant, amounting to 40 wt.% of total carotenoid in both uninfected G. pulex and G. roeseli. By contrast, we found only non-polar compounds with a predominance of esterified forms of astaxanthin in P. minutus larvae. No significant effect of infection on carotenoid content was evidenced in G. pulex and G. roeseli females. Our study highlights the use of a Matrix Solid Phase Dispersion as an efficient extraction method of both xanthophylls and non-polar pigments in small samples, including lipid-rich ones as P. minutus parasite. We discuss on the presumptive pathway leading to the formation of free astaxanthin in gammarids via hydroxy compounds, and on the accumulation of esters of astaxanthin in parasites.     

A Numerical Study of Local Variations in Tidal Regime of Tagus Estuary,Portugal

Tidal dynamics of shallow estuaries and lagoons is a complex matter that has attracted the attention of a large number of researchers over the last few decades. The main purpose of the present work is to study the intricate tidal dynamics of the Tagus estuary, which states as the largest estuary of the Iberian Peninsula and one of the most important wetlands in Portugal and Europe. Tagus has large areas of low depth and a remarkable geomorphology, both determining the complex propagation of tidal waves along the estuary of unknown manner. A non-linear two-dimensional vertically integrated hydrodynamic model was considered to be adequate to simulate its hydrodynamics and an application developed from the SIMSYS2D model was applied to study the tidal propagation along the estuary. The implementation and calibration of this model revealed its accuracy to predict tidal properties along the entire system. Several model runs enabled the analysis of the local variations in tidal dynamics, through the interpretation of amplitude and phase patterns of the main tidal constituents, tidal asymmetry, tidal ellipses, form factor and tidal dissipation. Results show that Tagus estuary tidal dynamics is extremely dependent on an estuarine resonance mode for the semi-diurnal constituents that induce important tidal characteristics. Besides, the estuarine coastline features and topography determines the changes in tidal propagation along the estuary, which therefore result essentially from a balance between convergence/divergence and friction and advection effects, besides the resonance effects.     

Comparison of Solea solea macroparasites between two nursery-continental shelf systems in the Bay of Biscay and the Portuguese coast

Digenean metacercariae of 0 year group common sole Solea solea ( n = 70) were more abundant in the embayed nursery of the Pertuis Charentais than in the Tagus estuary nursery. Macroparasite assemblages of adult sole ( n = 119) displayed only one species in common between the Bay of Biscay and the Portuguese coast continental shelves. These data highlighted the potential use of macroparasites as biological tags in various aspects of common sole ecology.     

Patterns of intermediate host use and levels of association between two conflicting manipulative parasites.

For many parasites with complex life cycles, manipulation of intermediate host phenotypes is often regarded as an adaptation to increase the probability of successful transmission. This phenomenon creates opportunities for either synergistic or conflicting interests between different parasite species sharing the same intermediate host. When more than one manipulative parasite infect the same intermediate host, but differ in their definitive host, selection should favour the establishment of a negative association between these manipulators. Both Polymorphus minutus and Pomphorhynchus laevis exploit the amphipod Gammarus pulex as intermediate host but differ markedly in their final host, a fish for P. laevis and a bird for P. minutus. The pattern of host use by these two conflicting manipulative parasites was studied. Their incidence and intensity of infection and their distribution among G. pulex were first examined by analysing three large samples of gammarids collected from the river Tille, Eastern France. Both parasites had low prevalence in the host population. However, temporal fluctuation in the level of parasitic infection was observed. Overall, prevalence of both parasite species was higher in male than in female G. pulex. We then assessed the degree of association between the two parasites among their intermediate hosts, using two different methods: a host-centred measure and a parasite-centred measure. Both measures gave similar results; showing random association between the two acanthocephalan species in their intermediate hosts. We discuss our results in relation to the selective forces and ecological constraints that may determine the pattern of association between conflicting manipulative parasites.     

A new goby, Glossogobius minutus (Teleostei: Gobiidae) from the south-west coast of India

A new species of gobiid fish Glossogobius minutus collected from the south-west coast of India is described. The morphological, morphometric and meristic characters were studied with special emphasis on osteology. The various characters are compared with those of G. giuris (Harn.) and G. biocellatus (Val.), the only other species of the genus reported from this region. Glossogobius minutus is a benthic omnivore and shows distinct sexual dimorphism.