Impact of predation on the polychaete Hediste diversicolor in estuarine intertidal flats

In estuarine sediment flats benthic macroinvertebrates are intensively consumed by a variety of predators, such as aquatic birds and nekton (mostly fish and crustaceans). However, there is still a lack of conclusive studies that evaluate if this predation has a relevant impact on the populations of those invertebrates, which are a key element of the estuarine food chain. In the Tagus estuary we experimentally tested and quantified the impact of predation on the polychaete Hediste diversicolor, one of the most important prey for a variety of predators in many estuaries. Using an exclusion experiment, we compared the seasonal variation in the densities of H. diversicolor from February to November in sediment plots (1) available to both bird and nekton predators, (2) just to nekton, and (3) without predators. We also followed changes in the abundance of potential predators throughout the study. The lowest densities were systematically observed in the plots accessible to all predators, followed by those which excluded just birds, and finally by those that excluded all predators. The exclosures were in place for 9 months, at the end of which the average density of H. diversicolor in the plots protected from all predators was eight times greater than in those without any protection. These results demonstrate that predation had a major impact on the densities of H. diversicolor. The relative importance of bird and nekton predation varied along the study, and this seems to be determined by different peaks of abundance of the two types of predators. However, when present in high densities, birds and nekton seem to have a similar impact on H. diversicolor. Our results suggest that predation is a key factor on the population dynamics of H. diversicolor. In addition, the levels of predation that we observed suggest that this polychaete can be a limited resource, and this could have major ecological consequences for predators for which it is a key prey.     

Evidence of estuarine nursery origin of five coastal fish species along the Portuguese coast through otolith elemental fingerprints

Connectivity is a critical property of marine populations, particularly for species with segregated juvenile and adult habitats. Knowledge of this link is fundamental in understanding population structure and dynamics. Young adults of commercially important fish species Solea solea, Solea senegalensis, Platichthys flesus, Diplodus vulgaris and Dicentrarchus labrax were sampled off the Portuguese coast in order to establish preliminary evidence of estuarine nursery origins through otolith elemental fingerprints. Concentrations of Li, Na, Mg, K, Mn, Cu, Zn, Sr, Ba and Pb in the otolith section corresponding to juvenile's nursery life period were determined through laser ablation inductively coupled plasma mass spectrometry. Element: Ca ratios in coastal fish differed significantly amongst collection areas, except for Platichthys flesus, and were compared with the elemental fingerprints previously defined for age 0 juveniles in the main estuarine nurseries of the Portuguese coast. Identification of nursery estuaries was achieved for four of the species. Assigned nursery origins varied amongst species and differences in the spatial scale of fish dispersal were also found. Diplodus vulgaris was not reliably assigned to any of the defined nurseries. Overall, results give evidence of the applicability of estuarine habitat tags in future assessments of estuarine nursery role. Research developments on the links between juvenile and adult habitats should contribute for the integrated management and conservation of nurseries and coastal stocks.     

The chemical composition of specular hematite from Tilkerode, Harz, Germany: implications for the genesis of hydrothermal hematite and comparison with the Quadrilátero Ferrífero of Minas Gerais, Brazil

The black shale-hosted selenide vein-type deposit at Tilkerode, eastern Harz, Germany, has specular hematite enclosed in clausthalite (PbSe). The specular hematite has Ti and V in amounts of up to ～1 wt.% TiO₂ and ～3 wt.% V₂O₅, and subordinate, but important, contents of Mo (22–372 ppm) and B (up to 68 ppm). The Tilkerode hematite serves as a reference for hydrothermal hematite formed at relatively low temperatures (<150 °C). The composition of the Tilkerode hematite is compared with that of two generations of specular hematite from itabirite-hosted iron-ore deposits in the Quadrilátero Ferrífero of Minas Gerais, Brazil. The first generation of specular hematite represents an early tectonic hematitisation of dolomitic itabirite at Águas Claras and occurs as fine-grained crystals. Reconnaissance data indicate that the Águas Claras hematite is poorer in Ti and V, relative to the Tilkerode hematite, but has ～5–10 ppm B and ～7–11 ppm Li. The second generation of specular hematite defines the pervasive tectonic foliation of the Gongo Soco iron ore. This hematite has Ti contents of up to ～2 wt.% TiO₂ and subordinate amounts of V (62–367 ppm); its B and Li concentrations are mostly below <2 ppm B and <1 ppm Li. The presence of Ti and B in the Tilkerode hematite can be explained by highly saline, B-bearing fluids that were capable of mobilising otherwise immobile Ti. The Mo signature of the Tilkerode hematite suggests that Mo was derived from the host black shale. In Minas Gerais, B and Li were incorporated into the early tectonic hematite from saline fluids at relatively low temperatures (Águas Claras) and then released during metamorphic hematite growth at higher temperatures, as suggested by the foliation-defining hematite without B–Li signature (Gongo Soco).     

Nursery fidelity,food web interactions and primary sources of nutrition of the juveniles of Solea solea and S. senegalensis in the Tagus estuary (Portugal): A stable isotope approach

Stable carbon and nitrogen isotopes were used to assess site fidelity of Solea solea and Solea senegalensis juveniles, to investigate food web interactions and to determine the dominant nutrient pathways in two nursery areas in the Tagus estuary, Portugal. Samples of water from the main sources and from the nursery areas and respective saltmarsh creeks were collected for isotope analysis, as well as sediment, benthic microalgae, saltmarsh halophytes, S. solea, S. senegalensis and its main prey, Nereis diversicolor, Scrobicularia plana and Corophium spp. While site fidelity was high in 0-group juveniles, it was lower for 1-group juveniles, possibly due to an increase in mobility and energy demands with increasing size. Analysis of the food web revealed a complex net of relations. Particulate organic matter from the freshwater sources, from each nursery's waters and saltmarsh creeks presented similar isotopic composition. Sediment isotopic composition and saltmarsh halophytes also did not differentiate the two areas. All components of the food web from the benthic microalgae upwards were isotopically different between the nursery areas. These components were always more enriched in δ¹³C and δ¹⁵N at the lower nursery area than at the nursery located upstream, appearing as if there were two parallel trophic chains with little trophic interaction between each other. A mixture of carbon and nitrogen sources is probably being incorporated into the food web. The lower nursery area is more dependent upon an isotopically enriched energy pathway, composed of marine particulate organic matter, marine benthic microalgae and detritus of the C₄ saltmarsh halophyte Spartina maritima. The two nursery areas present a different level of dependence upon the freshwater and marine energy pathways, due to hydrological features, which should be taken into account for S. solea and S. senegalensis fisheries and habitat management.     

Prey consumption by the juvenile soles, Solea solea and Solea senegalensis, in the Tagus estuary,Portugal

The soles Solea solea and Solea senegalensis are marine flatfish that use coastal and estuarine nursery grounds, which generally present high food availability, refuge from predators and favourable conditions for rapid growth. Two important nursery grounds for these species juveniles have been identified in the Tagus estuary, one in the upper part of the estuary (nursery A) and another in the south bank (nursery B). While S. solea is only present at the uppermost nursery area, S. senegalensis is present at both nurseries. Although they are among the most important predators in these nursery grounds, there are no estimates on their food consumption or on the carrying capacity of the system for soles. The Elliott and Persson [1978. The estimation of daily rates of food consumption for fish. Journal of Animal Ecology 47, 977–993] model was used to estimate food consumption of both species juveniles in both nursery areas, taking into account gastric evacuation rates (previously determined) and 24 h sampling surveys, based on beam-trawl catches carried out every 3 h, in the summer of 1995. Monthly beam trawls were performed to determine sole densities over the summer. Density estimates and daily food consumption values were used to calculate total consumption over the summer period. Sediment samples were taken for the estimation of prey densities and total biomass in the nursery areas. Daily food consumption was lower for S. solea (0.030 g wet weight d⁻¹) than for S. senegalensis (0.075 g wet weight d⁻¹). It was concluded that thermal stress may be an important factor hindering S. solea's food consumption in the warmer months. Total consumption of S. solea over the summer (90 days) was estimated to be 97 kg (wet weight). Solea senegalensis total consumption in nursery A was estimated to be 103 kg, while in nursery B it was 528 kg. Total prey biomass estimated for nursery A was 300 tonnes, while for nursery B it was 58 tonnes. This suggests that food is not a limiting factor for sole in the Tagus estuary. However it was concluded that more in-depth studies into the food consumption of other species and prey availability are needed in order to determine the carrying capacity of this system for sole juveniles.     

Estuarine production of resident and nursery fish species: Conditioning by drought events?

The production of resident (Pomatoschistus minutus and Pomatoschistus microps) and marine juvenile fish species using the Mondego estuary (central Portugal) as nursery grounds (Dicentrarchus labrax, Platichthys flesus, Solea solea), was assessed in order to: (1) understand the potential of the estuary for fish production; (2) know the production of nursery fish species likely to be exported to the coastal stocks; and (3) how anthropogenic and natural stress could influence the estimated production. Sampling occurred from June 2003 to May 2006 and together the 5 species in study comprised around 70% of the whole fish community numbers and biomass. Increasing drought conditions were observed, starting with a normal hydrological year in 2003 until attaining a severe drought in 2005, which resulted in low river discharges (1/3 of the mean river discharges in 2003). Additionally, high water temperatures were observed in 2003 and 2005 (24 and 26 °C, night temperatures). The secondary production was estimated using the increment summation method, after recognition of the cohorts. Production was in general lower in the Mondego estuary when compared to other systems, which was associated to the estuary's small area (only 3.4 km², less than 1/4 of area compared to other studied systems). Dicentrarchus labrax was among the most productive species. Production decreased in the drought year for all species, especially evident for D. labrax, P. minutus and P. flesus. No direct effects could be attributable to the salinity and temperature variations and to the low freshwater discharges (resulting from the drought and high temperatures), yet these were pointed as probable major reasons for the decreased production. A significant reduction (15–45% reduction in the estuarine production) was also concluded for the potential production to be exported for coastal areas by the nursery species in the drought conditions.     

Nursery use patterns of commercially important marine fish species in estuarine systems along the Portuguese coast

Analysing the estuarine use patterns of juveniles of marine migrant fish species is vital for identifying important sites for juveniles as well as the basic environmental features that characterize these sites for different species. This is a key aspect towards understanding nursery function. Various estuarine systems along the Portuguese coast (Minho, Douro, Ria de Aveiro, Mondego, Tejo, Sado, Mira, Ria Formosa and Guadiana) were sampled during Spring and Summer 2005 and 2006. Juveniles of commercially important marine fish species Solea solea, Solea senegalensis, Platichthys flesus, Diplodus vulgaris and Dicentrarchus labrax, predominantly 0-group individuals, were amongst the most abundant species and had distinct patterns of estuarine use as well as conspicuous associations with several environmental features. Juvenile occurrence and density varied amongst estuaries and sites within them, and differed with species. Sites with consistently high juvenile densities were identified as important juvenile sites (i.e. putative nursery grounds). Through generalized linear models (GLM), intra-estuarine variation in occurrence and density of each of the individual species was largely explained by environmental variables (temperature; salinity; depth; percentage of mud in the sediment; presence of seagrass; importance of intertidal areas; relative distance to estuary mouth; macrozoobenthos densities; and latitude). Decisive environmental factors defining important sites for juveniles varied depending on the system as a result of different environmental gradients, though there were common dominant features for each species regardless of the estuary considered. Analysed environmental variables in the GLM also accounted for inter-estuarine variation in species' occurrence and density. In several estuaries, the identified important juvenile sites were used by many of these species simultaneously and may be of increased value to both management and conservation. Overall, the variability in site features amongst estuaries highlighted the tolerance of these species to different available environmental conditions and provided fundamental information for future spatially explicit modelling of their distribution. This should ultimately enable the prediction of species response to habitat alterations.     

Comparing quality of estuarine and nearshore intertidal habitats for Carcinus maenas

Estuarine and nearshore marine areas are vital habitats for several fish and benthic invertebrates. The shore crab Carcinus maenas (Crustacea: Brachyura: Portunidae) inhabits a variety of coastal, estuarine and lagoon habitats. At low tide, habitat structural complexity may be most important for crabs in the intertidal, providing refuge from predation and desiccation. The quality of different vegetated and nonvegetated estuarine and rocky shore habitats in SW Portugal and SW England was evaluated for intertidal C. maenas populations. We estimated population density, size–structure, and potential growth (RNA/DNA ratios) to investigate habitat quality. Vegetated estuarine habitats supported higher crab densities, than nonvegetated estuarine and rocky shore habitats. Investigation of population size–structure revealed that all habitats seem important recruitment and nursery areas although estuarine habitats in SW Portugal appeared to support higher densities of new recruits than equivalent habitats in SW England. Significant variation was found in RNA/DNA ratios among habitats. Ratios were highest in the rocky shore suggesting a high quality habitat where growth potential is high. We speculate that competition from other top-predators (Pachygrapsus spp.) rather than low habitat quality may limit the occurrence of C. maenas in intertidal rocky shore habitats in SW Portugal. In estuarine environments RNA/DNA ratios were significantly higher in the vegetated than in the nonvegetated estuarine habitats in SW Portugal but not in SW England, suggesting geographic differences in the extent to which highly structure habitats represent high quality. Our results challenge the current paradigm that structured habitats are necessarily those of higher quality for C. maenas.     

Fish assemblages of small estuaries of the Portuguese coast: A functional approach

The importance of estuaries for coastal environments is widely acknowledged but their role, structure and ecological status have been the focus of recent scientific efforts mainly concerning large estuarine areas. In this work we used fish assemblages to establish, for the first time, the functional and ecological role of five small estuarine systems along the Portuguese south and southwest coasts. Our results showed that, at a functional approach, fish communities did not differ between estuaries, and that all systems presented a seasonal pattern in diversity values, ecological and feeding guilds. These small estuaries contribute to the support of coastal fish populations by providing temporary habitats to the critical life stages of marine species, shelter and feeding grounds, and should be considered in an ecological and conservation perspective.     

Nekton migration and feeding location in a coastal area – A stable isotope approach

Stable isotope analysis was used to investigate nekton movements and feeding location in a coastal area adjacent to a major European river, the Tagus, Portugal. Particulate organic matter isotopic signatures presented a gradient from the river towards the sea. Phytoplankton, zooplankton, polychaetes and the crab, Polybius henslowii, provided evidence of the incorporation of terrestrial organic matter into the lower levels of the food web, reflecting local isotopic signatures. Two fish species reflected the coastal isotopic gradient in δ¹³C, Diplodus vulgaris and Arnoglossus imperialis and the latter also presented isotopic differerences among the sites for δ¹⁵N. Alloteuthis subulata, Trisopterus luscus and Callionymus lyra were isotopicaly distinct among sites for δ¹⁵N. An increase of δ¹⁵N with length was detected for T. luscus and C. lyra, possibly showing ontogenic trophic level changes. Since A. subulata did not present differences in length and still showed isotopic distinction for δ¹⁵N, among areas, it was concluded that local biogeochemical factors may also have an influence. Diplodus bellottii and Dicologlossa cuneata did not reflect any isotopic signature reflecting their wide migration and feeding across the coastal area. Central isotopic ranges, defined as the site mean values for δ¹³C and δ¹⁵N ± 1‰ were determined for each species and site and those deviating from these were considered transient individuals. Central isotopic ranges accounted for 87% of A. imperialis, 80% of A. subulata, 77% of T. luscus, 67% of C. lyra and 50% of D. vulgaris. The number of individuals within each central isotopic range was surprisingly high for an open coastal area and comparable to those of more structured environments.