Growth parameters of deep-water decapod crustaceans in the Northwestern Mediterranean Sea: a comparative approach

 Relative and absolute growth were studied in 17 species of deep-water decapod crustaceans, spanning nine families of six different infra-orders, in the Northwestern Mediterranean Sea. The overall maximum abundance of these species lay between 200 m and 750 m (i.e. upper- and mid-slope species). Relative and absolute growth rates were compared by contrasting the slopes of the size–weight relationships for the different species and calculating the von Bertalanffy growth-equation parameters asymptotic length (L _∞ ) and growth rate (k). The size–weight relationships differed significantly as function of the species' life habits. The results revealed a significant decrease in weight relative to size in mesopelagic species (which carry out diel vertical migrations), an almost isometric relationship between size and weight in the less mobile nektobenthic species, and a significant increase in weight relative to size in strictly benthic species. The mean allometric coefficient for each group increased significantly from mesopelagic to benthic species. However, no general trend was observed in the growth-performance index, Φ (an index used to compare absolute growth rates between species, as a function of habit and depth of maximum abundance for all species combined), suggesting that the deep-water decapod crustaceans studied have similar absolute growth rates. Nevertheless, comparison of growth-parameter and growth-performance index values within families did reveal differences. Mesopelagic species of the families Sergestoidae and Pasiphaeidae showed slightly increased growth rates with increasing depth of distribution. Nektobenthic species of the genus Plesionika followed a trend opposite to that shown by mesopelagic species, with a higher growth rate for the shallowest-dwelling species (P. heterocarpus) than the deepest-dwelling species (P. acanthonotus). Taking growth as one of the major components of an organism's energy budget, the growth rates for the decapod crustacean species in this study were significantly lower than those reported in the literature for shallow-water penaeid crustacean species (which are distributed in higher-temperature habitats than deep-water Mediterranean crustaceans) and higher than those reported for mesopelagic myctophid fish species. Hence, the well-defined growth trends shown by deep-water decapod crustacean species in the Northwestern Mediterranean Sea, compared to the less well-defined trends in the other taxa, is discussed in the framework of the overall dynamics of their ecosystem. Received: 25 May 1998 / Accepted: 27 September 1999     

Analysis of the structure of decapod crustacean assemblages off the Catalan coast (North-West Mediterranean)

We sampled the communities of decapod crustaceans inhabiting the depth zone between 3 and 871 m off the Catalan coast (North-West Mediterranean) from June 1981 to June 1983. The 185 samples comprised 90 species differing widely in their depth distributions. Multivariate analysis revealed four distinct faunistic assemblages, (1) littoral communities over sandy bottoms, (2) shelf communities over terrigenous muds, (3) upper-slope communitics, and (4) lower-slope or bathyal communities. The brachyuran crab Liocarcinus depurator is the most abundant species of the shelf assemblage, although L. vernalis dominates over the shallow sandy bottoms of the shelf. The dominant species of the upper-slope assemblage are nektobenthic species (Solenocera membranacea, Plesionika heterocarpus, Processa canaliculata), pelagic species (Pasiphaea sivado, Sergestes arcticus), and benthic species (Macropipus tuberculatus, Munida intermedia, Nephrops norvegicus). Aristeus antennatus comprise most of the biomass of the lower-slope community, which supports a greater diversity than the other assemblages. The main assemblages appear to be related to different hydrological characteristics, the extent of seasonal fluctuations, and to the changes in sediment structure associated with changes in the steepness of the bottom.     

Biological patterns and near-bottom population characteristics of two pasiphaeid decapod crustacean species, Pasiphaea sivado and P. multidentata, in the north-western Mediterranean Sea

Biological aspects of the only two epi-/mesopelagic pasiphaeid species distributed along the continental marginof the north-western Mediterranean Sea were studied over the period 1991-1994. The annual reproductive cycle, reproductive output, sex and size distributions by depth (near-bottom distribution), and growth were analysed using 4156 specimens of Pasiphaea sivado (Risso, 1816) and 5491 specimens of P. multidentata Esmark, 1866. The reproductive cycle in the shallower-dwelling species, P. sivado, was nearly continuous, in contrast to the reproductive cycle in P. multidentata, which was markedly seasonal in nature. Both species exhibited similar size and population structure patterns with depth, with individual size increasing along the bottom depth gradient. The shallower-dwelling species had a significantly higher reproductive output level, in terms of the relative number of eggs, than the deeper-dwelling species. Biological trends for each of the species have been compared. The comparative results for these two congeneric species revealed that the biological parameters analysed were closely linked to their population structure. The slight differences in the depth range inhabited by each species have been considered the main environmental factor affecting the life histories of these two species. The data presented supported the hypothesis that depth exerted an important influence on the biological patterns of deep-water species, with the seasonality of reproductive processes increasing in deeper-dwelling species.     

Subcellular distribution of zinc and cadmium in the hepatopancreas and gills of the decapod crustacean Penaeus indicus

The decapod crustacean Penaeus indicus accumulated Cd and Zn in different subcellular compartments of hepatopancreas and gill cells. Most of the Cd and part of the Zn accumulates within the soluble fraction of the cells, while the remainder of the Zn is found in insoluble inclusions, associated with P, Ca, Mg and Si in B-, F- and R-cells in the hepatopancreas, and haemocytes, nephrocytes and epithelial cells in the gills. No presence of Cd was observed in metal-rich inclusions in any cell analysed. Metallothionein-like proteins (MTLP), analysed by differential pulse polarography, were present in the hepatopancreas (12–18 mg g⁻¹) and gills (7–8 mg g⁻¹) of metal-exposed prawns. Binding to MTLP is the detoxification mechanism for cadmium, while the detoxification of zinc involves both binding to MTLP and incorporation into insoluble metal-rich inclusions.     

A morphological approach for relating decapod crustacean cephalothorax shape with distribution in the water column

Cephalothorax profiles for individuals of 18 decapod crustacean species were obtained from pictures of specimens captured using various experimental bottom trawls in the Mediterranean basin, at depths ranging from 200 to 4000 m in 2000 and 2002. Profiles were compared using the shape (outline) of the entire cephalothorax (including and excluding the rostrum). Principal component analysis (PCA) of the profiles, rostrum included, yielded two large species groupings related to rostrum size, species having a long rostrum being clustered on the positive portion of the first axis and species having a short rostrum being clustered on the negative portion of this same axis (the F1 axis explained 72.17% of the total variance). The PCA separated pelagic and endobenthic animals (with short rostrums) from nektobenthic animals (with long rostrums). Only the two deepest-dwelling species (dwelling at depths below 2000 m) were distanced from their respective groups. The pelagic shrimp Acanthephyra pelagica was clearly grouped with the nektobenthic species despite being reported to have a pelagic habitat. The nektobenthic deep-sea species Nematocarcinus exilis has a relatively medium-size rostrum and was placed among the pelagic–endobenthic species. The endobenthic and pelagic groups remained stable even when rostral morphometrics were excluded from the PCA analysis. The hypothesis that predation and diet are the fundamental determining factors for all habitats throughout most of the lifetimes of individuals and hence that species will evolve specialised phenotypes to adapt to local environmental conditions is considered in the discussion.Communicated by O. Kinne, Oldendorf/Luhe     

Megabenthic decapod crustacean assemblages on the Galician continental shelf and upper slope (north-west Spain)

The structure of megabenthic decapod crustacean assemblages on the Galician (north-west Spain) continental shelf (100 to 200 depth) and upper slope (200 to 500 m) was analyzed based on surveys carried out in autumn and spring, from 1980 to 1987. Forty species belonging to 19 families were caught. The portunid crab Polybius henslowii, a species with pelagic phases, was the most abundant species, but displayed strong spatial and temporal fluctuations. Other dominant species were the Norway lobster Nephrops norvegicus, the portunid Liocarcinus depurator, the galatheids Munida intermedia and M. sarsi, and the shrimps Solenocera membranacea, Plesionika heterocarpus, Pasiphaea sivado and Dichelopandalus bonnieri. Total abundance and biomass (average values excluding Polybius henslowii = 255 individuals and 2.06 kg/30 min tow) and species richness and diversity, H′ (6.85 species and H′ = 1.45 per tow) displayed a significant positive correlation with depth, and strong interannual fluctuations. The factors determining community organization were depth and, to a lesser extent, spatial structure. There was clear evidence of bathymetric zonation, differentiating between species characteristic of the slope (D. bonnieri and Pasiphaea sivado), shelf-slope edge (Macropipus tuberculatus, Pontophilus spinosus, Munida sarsi, S. membranacea, Processa spp.) and shelf (L. depurator, Macropodia tenuirostris, Paguridae and Chlorotocus crassicornis). The spatial zonation was related to changes in oceanography and sediment along the continental margin. Goneplax rhomboides, N. norvegicus, C. crassicornis and Alpheus glaber are benthic species which generally exhibit burrowing behaviour, and they were found mainly in the southern area where there are fine sediments due to the outwelling from the Rías Baixas. Different benthopelagic shrimps (Pontophilus spinosus, Plesionika heterocarpus,Processa spp. and Pasiphaea sivado) were typical of the zone just north of Fisterra, characterized by a convergence of water masses bringing about an increase in productivity due to upwelling. The benthic anomuran and brachyuran crabs Munida intermedia, M.␣sarsi, L.␣depurator and Macropipus tuberculatus were characteristic of the northwestern zone between Fisterra and Estaca, where the infauna reaches high biomass despite coarser sediments with a lower concentration of organic material than in the southern area. Lastly, both the Paguridae and Macropodia tenuirostris were species typically found in the waters in the northern shelf. Based on interannual changes in assemblage structure, two periods could be distinguished: between 1980 and 1984, when Polybius henslowii, D. bonnieri and Pasiphaea sivado had abundance peaks; and another period from 1985 to 1987 when L. depurator, Munida intermedia, M.␣sarsi and Macropipus tuberculatus increased in abundance. Received: 21 May 1996 / Accepted: 9 August 1996     

Mediterranean octocorallian communities and the abiotic environment

Octocorallian communities (Coelenterata: Anthozoa) were investigated in 40 underwater stations near Banyuls-sur-Mer (France), as representatives of Mediterranean benthic fauna. The stations, with depths ranging from 0.3 to 40 m, were all sampled by means of SCUBA-diving. Along with a faunistic inventory, the main abiotic properties of each station were examined, such as irradiance, temperature, water movement, slope and presence of sediment. By means of cluster analysis and factor analysis, the main properties of which are briefly outlined in this paper, the existence of 5 separate communities is demonstrated. Two major abiotic factors emerge from this study, viz. submarine daylight and the presence or absence of sediment. Although community structures are found to be correlated with the physical environment, no causal relationships have been established. It is suggested that the tolerance of larval and young adult stages play an important role in this respect.     

Molt cycle alterations in behavior,feeding and diel rhythms of a decapod crustacean,the spiny lobster Panulirus argus

The crustacean molt cycle manifests extensive behavioral changes in addition to physiological and integumentary modifications. The paucity of quantitative studies led us to characterize molt stage dependent alterations in rhythmic locomotor, feeding, and agonistic behavior of subadult spiny lobster Panulirus argus held grouped and solitarily in simulated natural conditions. Non-disruptively determined molt stages were defined as proportions of intermolt duration. Significant nocturnal rhythmicity persisted through the full molt cycle, though daily form varied as a function of activity level and molt stage. A distinct early evening peak signaled initiation of foraging and walking behavior. Thereafter, rhythm amplitude either declined progressively (low activity: unimodal pattern), remained high (high activity: nocturnal plateau), or exhibited a secondary early morning peak (intermediate activity: bimodal and polymodal patterns). Activity ceased at or immediately prior to sunrise. Ecdysis was predominantly nocturnal, probably reflecting species specific spatiotemporal movement patterns and social behavior. A selective advantage of nocturnal ecdysis was postulated concerning avoidance of cannibalism and diurnal predators. Locomotor activity and feeding rates were not equivalent through the molt cycle, though both peaked in stages B₂-C₁. Locomotor activity remained high in metecdysis, decreased sharply at proecdysis initiation (D₀), reaching lowest levels in D₁-D₃, then increased in D₃ through B₁. Activity dropped steeply at ecdysis, though lobsters were capable of intense and coordinated activity. Feeding decreased slowly through metecdysis after the B₂-C₁ maximum, then increased temporarily in C₄ and D₀, indicating heightened feeding motivation. This contrasts with the locomotor activity decrease at proecdysis. Food consumption declined rapidly in D₁ and D₂ and ceased at the D₂-D₃ transition. Feeding resumed in B₁ or B₂, intensifying to maximum in late B₂. Feeding remained relatively constant within stages, whereas locomotor activity varied greatly, though both correlated with metabolic needs. Grouped and solitary lobsters displayed similar patterns of foraging and walking, equivalent to those of locomotor activity and feeding of solitary individuals. Frequency of agonistic interactions (not aggression per se) remained relatively constant through the cycle, peaking in metecdysis, though the highest relative proportion occurred near ecdysis. Lobsters then were submissive and avoided physical contact with conspecifics. Clearly, locomotor activity, feeding, and social behavior of P. argus are not simply determined. Indeed, behavior is distinctly phase coordinated with varying metabolic requirements dependent on the proximity to ecdysis and ecological pressures.     

Growth rates of the anchovyEngraulis encrasicolus and the sardineSardina pilchardus in the Northwestern Mediterranean Sea

The growth rates of the anchovyEngraulis encrasicolus Linnaeus, 1758 and the sardineSardina pilchardus Walbaum, 1792 in the Northwestern Mediterranean Sea between January 1987 and June 1989 was determined by means of otolith interpretation validated by length-frequency analysis. The growth-performance index was higher for the anchovy than for the sardine, in contrast to other areas where these species coexist, where the sardine generally displays a higher growth rate than the anchovy. The possible causes of this difference between geographical areas are discussed.     

Myosin ATPase activity in an estuarine decapod crustacean,Scylla serrata,as a function of salinity adaptation

The myosin ATPase activity of the flexor muscle of an estuarine crab, Scylla serrata, was studied in relation to salinity adaptation. The enzyme is activated more by calcium than by magnesium; it exhibits maximum activity at pH 9.0, and substrate inhibition above 0.5 mM ATP. The enzyme activity increases in crabs adapted to higher salinities. The enzyme from normal (70% sea water) crabs shows two pH optima; one at pH 7.0, the other at pH 9.0. The neutral optimum shifts to pH 6.0 upon adaptation to full strength sea water, but disappears upon adaptation to 25% sea water. The enzyme from normal crabs shows an optimum at 30 °C; adaptation to full strength sea water raises this optimum to 38 °C, whereas adaptation to 25% sea water decreases it to 24 °C. These changes are discussed in relation to estuarine conditions.     

Vertical distributions of decapod crustacean larvae and pelagic post-larvae over Great Sole Bank (Celtic Sea) in June 1983

Variations in the vertical distributions of pelagic stages of decapod crustaceans are described from five hauls taken over the Great Sole Bank in June 1983. Analyses of the results discriminated between the distributions mainly according to the time of day at which the hauls were taken, suggesting that diel variations were their most significant feature. Substantial ontogenetic variations were observed in the distributions of Processa canaliculata.     

Coastal hazard due to submarine canyons in active insular volcanoes: examples from Lipari Island (southern Tyrrhenian Sea)

The recent high-resolution multibeam bathymetry surveys around Lipari Island allowed to evidence several submarine canyons, whose head often cut back up to very shallow water and at a few tens of meters far from the coast. These canyons are mainly located in the eastern and southern side of the island and are characterized by an ongoing retrogressive (landward) erosion, that also controlled the shape and the evolution of the coastline. The canyon heads are formed by minor slide scars. By coupling slide scar morphometry and simple numerical model we have been able to roughly estimate the potential tsunami wave amplitudes generated by related slope failures. Moreover, the retrogressive erosion of canyon heads can be claimed as a cause of the enhanced subsidence reported in the last few thousand years in the eastern part of Lipari, where the main villages are located. Based on these evidence, we propose a first assessment of the coastal hazard due to marine retrogressive activity in the largest and most densely populated island of the Aeolian Archipelago.     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

Life history and fishery of Lepidopus caudatus (Pisces: Trichiuridae) in the Catalan Sea (Northwestern Mediterranean)

The frostfish Lepidopus caudatus (Euphrasen, 1788) is a mesopelagic species; it occurs along the shelf and slope down to 450 m in the Catalan Sea. Its' fishery, population structure, growth, diet and reproduction was studied on the basis of commercial statistics from 1984 to 1990 and on samplings made in 1988 and 1989. L. caudatus is fished by two types of commercial method, trawl and long-line; catches by the latter method display a marked seasonality. The minimum size of fish caught is 26 cm total length by trawling and 71 cm by long-line. Maximum sizes are similar for both gears: 196 and 188 cm, respectively. L. caudatus exhibits fast growth and attains an age of 8 yr. It feeds on small mesopelagic prey. It is a partial spawner reproductive activity occurring from April to November. Males attain sexual maturity at 97 cm and females at 111 cm total length.     

Relationships between nucleic acid concentrations and primary production in the Catalan Sea (Northwestern Mediterranean)

We explored the relationships between classical estimators of autotrophic biomass and primary production, such as chlorophyll a concentration and ¹⁴C-fixation rates, and biochemical indices based on DNA and RNA determinations, which have been proposed as indicators of physiological state in natural plankton populations. The measurements were made during two cruises across the Catalan Front, carried out in May 1989 and February 1990, corresponding respectively, to periods of stratification and moderate mixing. DNA and RNA concentrations (measured by a double-staining fluorimetric technique) were significantly correlated with chlorophyll a in February 1990, but not in May 1989, when a marked deep chlorophyll maximum was present. Significant positive correlations between RNA concentration and primary production and between RNA: DNA and primary production were found during both surveys, probably reflecting both higher RNA concentrations per cell and enhanced bacterial and microheterotrophic growth in high primary production situations. The results support the potential usefulness, in biological oceanography, of biochemical indicators based on DNA and RNA concentrations.     

Horizontal,vertical and seasonal distributions of epiphytic hydrozoa on the algaHalimeda tuna in the Northwestern Mediterranean Sea

The thalli ofHalimeda tuna (Ellis & Solander) grow horizontally from rocky surfaces and present an upper and lower surface which can be colonized by epiphytes. Thalli ofH. tuna were collected twice a month from March 1985 to March 1986 at Tossa de Mar (northeast Spain). Thirty five species of hydroid were recorded living onH. tuna thalli; of these, the 15 most abundant species were classified into three groups on the basis of their horizontal zonation on the thalli: those most abundant on the proximal, medial or distal articles. Those hydroids common on the proximal (oldest) articles were relatively large and were present throughout the year, those common on the medial articles were smaller and often occurred in dense monospecific patches, while those common on the distal (youngest) articles were present for only short periods and were the most opportunistic hydroids found onH. tuna. We hypothesize that this zonation reflects interspecific niche-selection, enabling successful competition for space with other hydroids, algae and bryozonas. Three hydroid species were more abundant on the upper surfaces of the algae than on the lower, probably because of gravitationally enhanced settlement of planula larvae coupled with (unusually) limited hydrorhizal growth on the underside. Three opportunistic species were more common on the lower surfaces of the thalli, possibly reflecting the greater availability of substratum in the shade, where epiphytic algae were less abundant. Many hydroids declined in abundance in summer because of the death of old thalli, the growth of new thalli and of apical articles on existing thalli, and possibly because of interspecific competition with epiphytic algae. The death of old thalli and the growth of new thalli reduced the abundance of epiphytic algae; this may have facilitated the increase in abundance of hydroids recorded in the autumn.     

Physical and biological gradients in a submarine cave on the Western Mediterranean coast (north-east Spain)

Submarine caves are semi-closed subsystems within the marine littoral ecosystem. One of the most striking features is the progressive reduction of the number of phyla, species, and biomass towards the interior of such a cave. The decrease in biomass has been explained as the result of low exchange rates of the water inside the caves which limit the supply of oxygen and food particles from the exterior. Many authors have proposed that this reduced exchange rate is associated with the occurrence of marked gradients in temperature, salinity, oxygen and light. Measurement of faunal abundance of a cave in the littoral zone of the Medes Islands (Catalonia, north-east Spain) made between June 1983 and July 1984, revealed a zonation similar to that in other Mediterranean caves. Animal biomass decreased from 260 g ash-free dry weight m^-2 at the entrance to 80 g ash-free dry weight m^-2 at the end of the cave (50 m from the mouth). A special technique for remote water-sampling was developed to avoid problems due to turbulence caused by divers in this semi-closed system. This involved the installation of a series of plastic tubes at different points along the axis of the cave. The water was brought to the surface using a battery-powered electric pump. The reliability of this method was verified by comparing the results with samples collected directly by divers. Sampling both in winter and summer showed no longitudinal gradients for temperature, salinity, oxygen, chlorophylla and the number and volume of suspended particles. These results suggest that there is a constant circulation within the cave which guarantees water-exchange. The decrease in biomass cannot be explained by physico-chemical gradients due to insufficient exchange with the external system. On a finer scale, however, along the walls of the cave there may exist boundary layers that are not affected by the general circulation. This could account for the observed decrease in the biomass of benthic organisms.     

The minimal area problem in invertebrate communities of Mediterranean rocky substrata

Different ways of obtaining information about minimal area are studied. The classical species-area curve lacks objectivity. Curves based on similarity indices are objective, in so far that a critical threshold can be defined, where one admits that the minimal area is reached. Comparison is made between Sørensen's qualitative similarity index (based on presence-absence) and Kulczynski's quantitative similarity index (based on numbers of individuals and/or percent cover). In the shallowwater rocky habitats studied, two types of community were distinguished. The first, miniaturized communities are found in dark caves. Their minimal area is about 4,000 cm². The second, coarser, communities occur on open rocks, either horizontal, sloping or vertical. Their minimal area is 20,000 cm². Calculations were based mainly on Octocorallia (Coelenterata) which are assumed to be characteristic of the community as a whole.