Ecological understanding for fishery management: Condition and growth of anchovy late larvae during different seasons in the Northwestern Mediterranean

The fishery of the European anchovy Engraulis encrasicolus in the Mediterranean needs several ecological approaches to be properly managed. As such, several surveys were carried out to study the ecology of larvae and juveniles of this species, which reproduces during the warmest period of the year (May through September) in the Gulf of Lions. In particular, we studied the late larvae (15 mm total length until metamorphosis), especially as other authors have focused on larvae below that size. Unexpectedly, we also collected anchovy late larvae during the December 2007 survey, whose range in size corresponded to a later spawning period than previously reported. Differences in the nutritional condition of these larvae were assessed by comparing indices of lipid composition and estimating growth rates from otolith measurements to provide information on the probability of survival between the two groups. The analysis of fatty acids, used as tracers of trophic relationships, indicates that these larvae fed mainly on zooplankton. Nutritional conditions of summer and late autumn larvae were very similar. In contrast, growth rates were higher for August larvae, probably due to the different temperatures in the two seasons. Our results are especially relevant in an ecological context where the increasing mean water temperatures in the Western Mediterranean could favor the extension of the anchovy spawning period up to late-Autumn months.     

Interranual variability in horizontal patterns of larval fish assemblages in the northeastern Aegean Sea (eastern Mediterranean) during early summer

Larval fish community structure was studied in the northeastern Aegean Sea (NEA) over an area influenced by the advection of Black Sea water (BSW). Sampling was carried out in early summer during a period of 4 years (2003–2006). Taxonomic composition and abundance presented high variability in space that remained relatively constant among years. Tow depth and indicators of trophic conditions in the upper water column (i.e., zooplankton displacement volume, fluorescence) explained significantly the structure of larval assemblages during all surveys. The northern continental shelf (Thracian and Strymonikos shelf), where a large amount of enriched, low salinity BSW is retained, was dominated by larvae of epipelagic species, mainly anchovy (Engraulis encrasicolus). Interannual changes in horizontal extension of the BSW seemed to match closely observed changes in the distribution of anchovy larvae. Mesopelagic fish larvae were particularly abundant beyond the continental shelf (over the North Aegean Trough) where a strong frontal structure is created between the low salinity waters of BSW origin and the high salinity waters of the Aegean Sea. Larvae of certain mesopelagic species (e.g., Ceratoscopelus maderensis) may occasionally be transported inshore when the prevailing current meanders towards the coast or feeds anticyclonic gyres over the continental shelf.     

Description of the Phytoplanktonic Community of the Oligotrophic Waters of the SE Aegean Sea (Mediterranean)

Abstract. A seasonal sampling program of five stations off the Island of Rhodes (SE Aegean Sea) was carried out in 1983–1984. Temperature, salinity, Secchi disk transparency, P-PO₄, N-NO₃, N-NO₂, N-NH₃, Si-SiO₂, and chl a were measured and phytoplankton species recorded. Cell concentrations and chl a varied seasonally. with the highest values in summer (l.2 times 10⁴ -1^--¹ total mean cells; 0.13 mgam^-3 total mean chl a ) and the lowest in winter (2.3 times 10³, 1^--¹ total mean cells; 0.06 mg^.m-^- total mean chl a ). A variation in cell abundance among stations was also noted. Quantitative relationships among the recorded taxa showed that diatoms and dinoflagellates were richer in species composition (88 and 58 total species, respectively) than coccolithophores (8 species) and other flagellates (8 species). Comparison of phytoplankton samples from different depths and stations by cluster analysis showed an irregularity or discontinuity in species associations. The SE Aegean Sea was characterized as oligotrophic on the basis of the estimated nutrient and phytoplankton concentration levels.     

Spatial, temporal and ontogenetic variation in diet of anchovy (Engraulis encrasicolus) on the Algerian coast (SW Mediterranean)

The diet of anchovy Engraulis encrasicolus was studied in three regions (Béjaia, Bénisaf and Ghazaouet) along the Algerian coast. Ontogenetic, spatial and seasonal variations in anchovy diet were investigated using multivariate analyses and analysed in relation with sea surface temperature and chlorophyll-a. 46 prey taxa of varying size between 0.57 mm (Euterpina acutifrons) and 6.8 mm (fish larvae) were recorded. Whatever the season, the region or the fish size, anchovy is exclusively zooplanktivorous and copepods were the most present prey, constituting 87% by number of the prey taken and found in 98% of the anchovy stomachs examined. However, their occurrence and number varied according to the different areas, seasons and fish size. During its first year of life, anchovy feeds almost exclusively on copepods (mainly small and medium size prey). As anchovy grows, copepods are gradually substituted by large crustaceans such as decapods and amphipods. Hierarchical cluster analysis, analysis of similarities (ANOSIM) and similarities percentage (SIMPER) indicated a distinct diet of anchovy of the bay of Bejaia from those of the bays of Bénisaf and Ghazaouet probably due to differences in hydrologic conditions. Diet differences also occurred between seasons. Summer and spring have distinct prey assemblages each and showed low diet similarities with the two other seasons. More prey species were found in the diet during winter (36) and autumn (30) and the vacuity index was lower in winter. Temporal variability in satellite-derived chlorophyll-a matched the seasonal variability in the diversity of the anchovy prey and feeding intensity as reflected by the vacuity index, suggesting further investigation of the potential use of satellite-derived chlorophyll-a data as a proxy for anchovy feeding intensity.     

Spatial variability of meiofaunal communities at areas of contrasting depth and productivity in the Aegean Sea (NE Mediterranean)

The abundance and community structure of metazoan meiofauna were studied in deep-sea sediments from the north and south Aegean Sea (Eastern Mediterranean) in summer 1997 and spring 1998. The two areas varied in their surface primary productivity with the northern area being more productive. Meiofaunal densities displayed strong spatial variability while no temporal changes were observed. Total metazoan density ranged from 128 to 1251 ind./10 cm², with significantly higher values in the north. Meiofaunal biomass ranged from 27 to 391 μgC/10 cm² with higher values also in the north. At all stations nematodes dominated the community, comprising on average more than 91% of the total abundance. On a broad scale, meiofaunal densities displayed a positive correlation with food availability (sediment-bound chloroplastic pigments, carbohydrates and lipids); carbon mineralization (an indicator of organic matter turnover) was significantly higher in the northern Aegean, providing evidence of high organic-matter input and intense benthic-pelagic coupling. The spatial structure of the nematode community indicated that the two areas were similar in terms of their dominant genera (Halalaimus, Acantholaimus, and Thalassomonhystera).     

Deep-water stands of Cystoseira zosteroides C. Agardh (Fucales,Ochrophyta) in the Northwestern Mediterranean: Insights into assemblage structure and population dynamics

Populations dominated by Cystoseira zosteroides, an endemic and threatened Mediterranean seaweed, colonize deep-water rocky habitats down to more than 50 m depth. Assemblages dominated by this species display high algal and invertebrate species richness. Algal biomass averages 1134 g dw m⁻². Erect and turf algae account for only 25% of total algal dry weight, while encrusting corallines are responsible for the remaining 75%. Sponges, bryozoans and ascidians constitute the dominant sessile macrofauna. Cystoseira zosteroides is the dominant erect algae, with a mean biomass of 60.6 g dw m⁻², and densities ranging from 4 to 7 plants m⁻². The alien turf alga Womersleyella setacea has a biomass of 104.2 g dw m⁻² and covers most of the understory substrate. The size-frequency distribution of C. zosteroides populations shows differences over time. Mean annual growth of the main axis is around 0.5 cm and mean annual mortality rate is lower than 2%. Recruitment was almost nil during the studied period of time (10 years). Processes structuring these deep-water Cystoseira stands must be driven by episodic disturbances, after-disturbance recruitment pulses, and long periods of steady growth that last at least 10 years. However, it is also possible that recruitment is irreversibly inhibited by the alien alga W. setacea in which case these old-growth stands are faced with extinction. The highly diversified assemblages and the low growth and low mortality rates of C. zosteroides indicate high vulnerability to natural and anthropogenic disturbances, and call for effective measures to ensure their conservation.     

Nursery areas of red mullet (Mullus barbatus), hake (Merluccius merluccius) and deep-water rose shrimp (Parapenaeus longirostris) in the Eastern-Central Mediterranean Sea

The spatial pattern of the nursery areas of red mullet (Mullus barbatus), hake (Merluccius merluccius) (Linnaeus, 1758) and deep-water rose shrimp (Parapenaeus longirostris) (Lucas, 1846) was studied in the South Adriatic and North Ionian Seas (Eastern-Central Mediterranean) applying geostatistical techniques and data from time series trawl surveys conducted in the area. The analysed variables were: R (number of recruits/km²) and R/Tot (fraction of recruits on the total sampled population). The structural analysis showed a spatial pattern of both variables characterized by continuity on a small scale. Predictions of nursery area localization with probability of finding recruits at different threshold values were obtained through median indicator kriging. For the red mullet the nurseries were mainly identified in the South Adriatic Sea off the Gargano peninsula and between Molfetta and Monopoli within 50 m in depth. The main concentration of hake juveniles was found to be between 100 and 200 m in depth along the Gargano peninsula and between Otranto and Santa Maria di Leuca, where a nursery of deep-water rose shrimp was also detected. An overlapping depth, between 100 and 200 m, was identified for hake and deep-water rose shrimp nurseries. Protection of these areas through limitations of fishing pressure is discussed.     

Human influence on seagrass habitat fragmentation in NW Mediterranean Sea

Habitat fragmentation in meadows of Posidonia oceanica, the most important and abundant seagrass in the Mediterranean Sea, was investigated at a region-wide spatial scale using a synthetic ecological index, the Patchiness Index (PI). We tested the hypothesis that human impacts are the major factor responsible for habitat fragmentation in P. oceanica meadows contrasting fragmentation of meadows located in “anthropized” areas with that of meadows located in areas with low anthropization and considered as virtually “natural”. We also related fragmentation of meadow with the morphodynamic state of the submerged beach (i.e. distinctive types of beach produced by the topography, the wave climate and the sediment composition) in order to investigate the influence of one natural component on the seagrass meadow seascape. Results demonstrated that fragmentation in the P. oceanica meadows is strongly influenced by the human component, being lower in natural meadows than in anthropized ones, and that it is little influenced by the morphodynamic state of the coast. The use of landscape approaches to discriminate natural disturbance from human impacts that affect seagrass meadows is thus recommended for the proper management of coastal zones.     

Environmental control on early life stages of flatfishes in the Lima Estuary (NW Portugal)

Several flatfishes spawn in oceanic waters and pelagic larvae are transported inshore to settle in the nursery areas, usually estuaries, where they remain during their juvenile life. Nursery areas appear as extremely important habitats, not only for juveniles but also for the earlier planktonic larval fish. Yet, the majority of nursery studies tend to focus only on one development stage, missing an integrative approach of the entire early life that fishes spent within a nursery ground. Thus, the present study assessed the influence of environmental parameters on the dynamics of the larval and juvenile flatfishes, throughout their nursery life in the Lima Estuary. Between April 2002 and April 2004, fortnightly subsurface ichthyoplankton samples were collected and juveniles were collected from October 2003 until September 2005. Larval assemblages comprised nine flatfish species, while only six were observed among the juvenile assemblages. Solea senegalensis and Platichthys flesus were the most abundant species of both fractions of the Lima Estuary flatfishes. Larval flatfish assemblages varied seasonally, without relevant differences between lower and middle estuary. Platichthys flesus dominated the spring samples and summer and autumn periods were characterized by an increase of overall abundance and diversity of larval flatfishes, mainly S. senegalensis, associated with temperature increase and reduced river flow. On the contrary, during the winter abundance sharply decreased, as a consequence of higher river run-off that might compromised the immigration of incompetent marine larvae. Juvenile flatfishes were more abundant in the middle and upper areas of the estuary, but the species richness was higher near the river mouth. Sediment type, distance from the river mouth, salinity, temperature and dissolved oxygen were identified as the main environmental factors structuring the juvenile flatfish assemblages. Juveniles were spatially discrete, with the most abundant species S. senegalensis and P. flesus associated with the middle and upper estuary, while the remaining species were associated with the lower estuarine areas. The larval fraction exhibited distinct dynamics from the juvenile estuarine flatfish community. Larval flatfishes showed a strong seasonal structure mainly regulated by biological features as the spawning season and also by seasonal variations of water characteristics. On the other hand, juvenile flatfishes were markedly controlled by site specific characteristics such as sediments structure, distance from the river mouth and salinity regime. The present study emphasized the idea that the environmental control varies throughout the ontogenetic development, stressing the importance of integrating all the early life of a species in flatfish nursery studies.     

Phytopigments and fatty acids in the gut of the deposit-feeding heart urchin Echinocardium cordatum in the southern North Sea: Selective feeding and its contribution to the benthic carbon budget

As part of a broader study on benthic-pelagic coupling in the southern North Sea, specimens of the common heart urchin Echinocardium cordatum were sampled for analyses on phytopigments and fatty acids in their guts. Results were interpreted in the context of feeding and ecological functioning of the heart urchins in the benthic system. Ingestion selection factors for both component groups were relatively high, 5 to 9 for chlorophyll a and 9 to 130 for total fatty acids. The data point to at least partially different sources of the pigments and of the fatty acids. Next to algal detritus, small infauna relatively rich in fatty acids might be preferentially co-ingested with the detritus. Due to digestive breakdown and absorption, the concentrations of pigments and fatty acids were importantly decreased, indicating a rather high digestion efficiency for this subsurface deposit feeder, up to 80%. The results indicate that E. cordatum increases its energy acquisition by strong selectivity and a high digestive efficiency. Optimal foraging is likely to apply on deposit-feeding invertebrates in relatively food-rich coastal environments as much as it does in the food-poor deep-sea environment. Using chlorophyll a as a proxy for carbon, the contribution of the urchin population to the momentary benthic carbon budget was calculated at 7% to 42%.     

Environmental influences on temporal patterns of settlement in two littoral labrid fishes in the Mediterranean Sea

Correlations between daily fish settlement and environmental and physical parameters were examined during two consecutive years in two labrid species (Symphodus roissali and Symphodus ocellatus) in the NW Mediterranean Sea. Both these species have a short planktonic larval duration (12±0.3 and 9±0.4 days, respectively) and limited offshore larval dispersal. Time-series analysis was used to compare the daily settlement patterns with various environmental variables, and non-linear generalized additive models (GAM) were used to examine the relationship between environmental conditions averaged over larval period and daily settlement patterns. Settlement was not uniform and took place as pulses of different length. Although the analyses detected a few significant correlations between day-to-day patterns in environmental variables and settlement (e.g. solar radiation, atmospheric pressure, daytime or nocturnal wind components), these were not consistent among years. In S. ocellatus, however, wind components (daytime or nighttime) explained part of the variance in settlement intensity. The correlations (GAM outputs) observed between settlement patterns and environmental variables (averaged during the whole planktonic period) suggest that calm weather during the planktonic period usually increases settlement success.     

Revisiting Peruvian anchovy (Engraulis ringens) trophodynamics provides a new vision of the Humboldt Current system

The Peruvian anchovy or anchoveta (Engraulis ringens) forages on plankton and is a main prey for marine mammals, seabirds, fish, and fishers, and is therefore a key element of the food web in the Humboldt Current system (HCS). Here, we present results from the analysis of 21,203 anchoveta stomach contents sampled during 23 acoustic surveys over the period 1996–2003. Prey items were identified to the genus level, and the relative dietary importance of different prey was assessed by determination of their carbon content. Variability in stomach fullness was examined relative to the diel cycle, the distance from the coast, sea surface temperature, and latitude, using generalized additive models (GAMs). Whereas phytoplankton largely dominated anchoveta diets in terms of numerical abundance and comprised >99% of ingested prey items, the carbon content of prey items indicated that zooplankton was by far the most important dietary component, with euphausiids contributing 67.5% of dietary carbon followed by copepods (26.3%). Stomach fullness data showed that anchoveta feed mainly during daytime between 07h00 and 18h00, although night-time feeding also made a substantial contribution to total food consumption. Stomach fullness also varied with latitude, distance from the coast, and temperature, but with substantial variability indicating a high degree of plasticity in anchoveta feeding behaviour. The results suggest an ecological role for anchoveta that challenges current understanding of its position in the foodweb, the functioning of the HCS, and trophic models of the HCS.     

Spatio-temporal patterns of abundance,size structure and body condition of Atherina boyeri (Pisces: Atherinidae) in a small non-tidal Mediterranean lagoon

Sand smelt, Atherina boyeri, is a very abundant, widespread, locally harvested species, able to successfully survive and reproduce over a wide range of abiotic conditions and aquatic ecosystem types. In the North Sea and in the Mediterranean Sea transitional ecosystems its life history is well known, but very little information is available about the influence of intra-ecosystemic abiotic gradients on micro-scale patterns of distribution, body size and condition of individuals. In a small non-tidal Mediterranean lagoon, the cohorts' structure and evolution of sand smelt was described and the temporal and spatial variability of abundance, size structure and body condition was addressed. The study was carried out at the Acquatina lagoon (Lecce, Italy) where four stations, situated in two habitat types along a confinement gradient, were sampled twice per month for one year with fyke nets. The abundance of collected individuals was very high throughout the sampling period, with a peak in Summer and a constant decrease afterwards. Measured Standard Length ranged from 20 to 90 mm and the majority of individuals were estimated to be within 2 years old. The confined station resulted more abundantly colonized, especially by smaller size classes, and characterized by a narrower size frequency distribution. Also relative body condition showed significant spatial variations, with larger individuals having a better condition than smaller ones where optimal temperature and dissolved oxygen levels were available. Size-specific temporal patterns of body condition followed the seasonal climatic oscillations and the species' cycle of growth and reproduction, with larger individuals reaching higher fatness than smaller ones during Autumn and Winter, thus having more chances to survive and contribute to the reproductive effort. The results of the present study emphasised that, even at a very small scale, the presence of strong environmental gradients can drive movements, habitat preference and condition of individuals and thus promote an intra-ecosystemic spatial heterogeneity comparable to that observed between ecosystems at the land–ocean or geographical scale.     

Contaminants and habitat choice in the Baltic Sea: Behavioural experiments with the native species, Monoporeia affinis, and the invasive genus, Marenzelleria

The invasive polychaete genus, Marenzelleria and the native amphipod, Monoporeia affinis are food and habitat competitors in the Baltic Sea. Previous studies have shown that moderate densities of Marenzelleria can affect the behaviour of M. affinis. To examine the short-term interactive effects of interspecific habitat choice and environmental contaminants a series of habitat colonisation experiments were performed. The contaminants examined included harbor sediments and sediment spiked with the antifouling substances, Cu, Zn and Irgarol. Polychaetes and amphipods were exposed to contaminants in single-species and two-species experiments. In spiked-sediment experiments, M. affinis showed clear dose-dependent response. These experiments verified that behavioural response of M. affinis to different habitats is a sensitive method for testing toxicity under controlled conditions. In experiments with three different harbor sediments and reference sediment both species showed the lowest preference for the reference sediment. This sediment also had the lowest content of quality food, indicating that factors such as food quality and quantity may override the disturbing effects of contaminants in natural sediments. The presence of Marenzelleria spp. did not affect amphipod habitat choice, indicating no short-term effects, which implies that both species can co-exist provided sufficient food is available.     

Feeding ecology of Ammothella longipes (Arthropoda: Pycnogonida) in the Mediterranean Sea: A fatty acid biomarker approach

Fatty acid analysis has proved valuable in determining seasonal trophic links and the feeding behavior in organisms in which these diet and trophic links cannot be inferred from stomach content analyses. Seasonal variations in total free fatty acid content (TFFA) and fatty acid composition of seston (<250 μm), the brown macroalgae Stypocaulon spp., polychaetes (Nereididae) and the pycnogonid Ammothella longipes have been used to establish their trophic links, with particular focus on seasonality and feeding ecology of A. longipes. Samples were collected in a coastal environment (NW Mediterranean Sea) at 7–10 m depth, in five different periods (August and October 2008, February, June and September 2009). Seston and Stypocaulon spp. samples did not show significant seasonal variations in TFFA content, while nereids showed a significant variation. Analysis of fatty acid profile showed high similarities of fatty acid composition between seston and Stypocaulon spp. Nereids were closer to seston and Stypocaulon spp. than A. longipes, which seemed to follow a seasonal trend. The results of this study reveal that A. longipes may change its feeding behavior depending on the season and available food. This pycnogonid species appears to be carnivore during spring and early summer but seems to feed on detritus when availability of prey diminishes during winter. Notable high amounts of odd-chain fatty acids are found in summer–autumn for this species, which may come from bacteria acquired from the detrital diet or from de novo biosynthesis from propionate. The results obtained provide new and valuable data on the understudied feeding biology of pycnogonids in general, and contribute to the understanding of their functioning of Mediterranean shallow oligotrophic systems and their trophic links.