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.     

Reproductive cycle and minimal length at sexual maturity of Engraulis encrasicolus (L.) in the Zrmanja River estuary (Adriatic Sea, Croatia)

The reproductive cycle of anchovy, Engraulis encrasicolus (L.), was studied from monthly random samples of purse seine catches. A total of 1477 anchovy specimens were collected from January to December 2003 in the Zrmanja River estuary (Novigrad Sea). The analysis was based on the temporal evolution of gonadosomatic index, mass and stage of gonads. The total length of anchovy ranged from 4.5 to 14.5 cm and mass from 0.56 to 19.80 g. Sex ratio was slightly different from 1:1; the females were insignificantly predominated (♂/♀ = 0.99). The period of reproductive activity was from April to September coinciding with the most developed stages of gonads as well as with the highest gonad weights, and gonadosomatic indices. To estimate the length at maturity, a sub sample of 454 anchovy was taken from May to July (peak of anchovy spawning period). The length at which 50% of anchovy were mature (L₅₀) was calculated to be 8.2 cm. The length–weight relationship of anchovy was described by the expression: W = 3.51 × 10⁻³ L_T^3.211 (r² = 0.998). The relationships between total length–standard length and total length–fork length are L_T = 1.1405L_S + 0.2420 and L_T = 1.0425 L_F + 0.3944, respectively.     

Geographic structure of European anchovy: A nuclear-DNA study

Atlantic–Mediterranean anchovies were genetically characterized at two polymorphic nuclear loci (intron 6 of two creatine-kinase genes) and compared to reference Engraulis albidus and E. encrasicolus samples from the northern Western Mediterranean to provide new insights into their geographic structure. Northeastern Atlantic anchovy, represented by one sample from the Canary archipelago and one sample from the Alboran Sea, were genetically distinct from Mediterranean E. encrasicolus (Weir and Cockerham's ^θ = 0.027–0.311), indicating geographic isolation from either side of the Almería–Oran oceanographic front. Generally smaller genetic differences were evident among anchovy populations from different sub-basins in the Mediterranean (^θ = − 0.019–0.116), the genetic differences between Black Sea and Ionian Sea/Aegean Sea anchovies being the strongest (^θ = 0.002–0.116). There was no evidence of the presence of E. albidus in our samples outside Camargue (northern shore of the Western Mediterranean). However, a sample from the southern Western Mediterranean appeared to be genetically intermediate between E. albidus and Mediterranean E. encrasicolus, indicating possible hybridization. Anchovy from the Benguela current system off southern Africa possessed allele frequencies characteristic of E. albidus at one locus and Northeastern Atlantic anchovy at the other locus, suggesting past introgression.     

Summer larval fish assemblages in the Southern Tyrrhenian Sea (Western Mediterranean Sea)

The purpose of this paper was to study the community structure, in terms of species composition, abundance and spatial distribution, of fish larvae in a wide coastal area of Sicily facing the Southeastern Tyrrhenian Sea, extending for 2300 km² from Cape Cefalù to the west, to Cape Rasocolmo in the east. This study analyses how species are assembled in relation to an inshore–offshore gradient and also how environmental conditions, determined by surface circulation patterns occurring in the Central Mediterranean at the local scale, determine the distribution patterns. Samples from 39 stations were collected using a 60‐cm Bongo net during an ichthyoplanktonic survey carried out in June 2006. In all, 62 taxa, representing 32 families, were identified. Cyclothone braueri (59.6%), Engraulis encrasicolus (9.2%) Lampanyctus crocodilus (4.3%) and Lampanyctus pusillus (4.1%) were the most abundant species. The results showed that the highest abundance value (14830.6 fish larvae per 10 m² sea surface) was observed in the western part of the study area. MDS, SIMPER and CCA analyses revealed well defined groups of stations and assemblages of larvae in accordance with an inshore–offshore gradient. The results of this study could have implications for the management of marine resources because the investigated area has already been identified as a nursery area for many pelagic and coastal fishes and a natural habitat for many species of high commercial interest.     

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.     

Fate of the Black Sea Acartia clausi and Acartia tonsa (Copepoda) penetrating into the Marmara Sea through the Bosphorus

In October 2005 spatial distribution of live and dead Acartia clausi and Acartia tonsa was studied in the Black and Marmara Seas and near the Marmara Sea inlet of the Bosphorus, in order to understand their fate upon transportation between two seas. The morphometric characteristics in both species from all studied areas, and the decreased abundance of A. clausi and A. tonsa from the Black Sea towards the Marmara Sea indicate that the Marmara Sea Acartia populations are formed by recruitment from the Black Sea. We observed mass mortality of A. clausi in the Marmara Sea near the Prince Islands. The majority of carcasses (66% of total A. clausi numbers in the Marmara Sea) were found in the salinity gradient layer.     

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).     

绿鳍马面鲀早期生长发育与摄食特性的研究

在人工培育条件下,采用实验生态学方法,对绿鳍马面鲀(Thamnaconusmodestus)仔、稚、幼鱼生长、发育与摄食特性的变化进行了研究。结果表明,绿鳍马面鲀仔、稚、幼鱼的各生长指标可拟合为具有1～2个生长转折点的2～3段直线,其中全长、体长、体质量、肛前长、头长、吻长和体高可拟合为3段直线,眼径可拟合为2段直线。头长在第一转折点后的生长速率显著高于其他指标,除头长和体高在第二个生长转折点后生长速率减慢外,其余各项生长指标生长速度均逐渐加快。3～4日龄"开口期"前后的仔鱼,摄食发生率和饱食率均较低,随后逐渐升高。摄食发生率在12日龄后达到100%,饱食率在25日龄后达到100%(消化道充塞度为3～4级)。绿鳍马面鲀仔、稚、幼鱼均为白天摄食类型,白天摄食量占全天摄食量的71.6%以上。随日龄增长,鱼苗饱食时间呈下降趋势,消化时间则呈上升趋势。了解绿鳍马面鲀早期的生长发育与摄食特性,可为人工育苗管理提供参考,以促进鱼苗生长,提高成活率。     

Habitat preferences and life history of the red scorpion fish, Scorpaena notata, in the Mediterranean

Scorpaena notata is a small, sedentary scorpaenid species widely distributed in the Mediterranean and adjacent waters of the Atlantic. In the western Mediterranean it inhabits coastal continental shelf bottoms. In the Balearic Islands, these bottoms are characterised by the presence of the facies with red algae, including both Peyssonnelia and mäerl beds. These beds enhance the structural complexity, biodiversity and secondary production of the soft bottoms. Due to the oceanographic conditions of the Islands, the facies with red algae are especially rich in terms of biomass and algal coverage, and are widespread distributed between 40 and 90 m depth, where trawlers exploiting the continental shelf operate. The present work studies the biology of S. notata and its relationship with habitat characteristics. Special attention is focused on the aspects related to fish condition and growth as a tool to assess the importance of the facies with red algae for fish.     

Biodiversity of zoobenthic hard-substrate sublittoral communities in the Eastern Mediterranean (North Aegean Sea)

The spatial dispersion of zoobenthos from sublittoral hard substrate communities in the northern part of the Aegean Sea has been studied during summer 1997 and 1998. Material was collected by SCUBA diving, by totally scraping off five replicate quadrates (400 cm² each) at three depth levels (15, 30, 40 m) from six sites located in Chalkidiki peninsula, plus one in Kavala Gulf. The examination of the 19,343 living specimens collected revealed the presence of 314 species. Though the multivariate analyses showed high similarity between stations, the structure of this sciaphilic algal community seems to have an increased spatial heterogeneity. Four distinct facies were recorded in accordance with the occurrence of different algal forms, the degree of hard substrate inclination and the water clarity. A short review on the biodiversity of sublittoral communities in the Mediterranean revealed the affinity between the western and the eastern basin and also among the photophilic and the sciaphilic algal communities.     

Carbon, nitrogen and phosphorus stoichiometry in Japanese anchovy (Engraulis japonicus) from the Huanghai Sea, China

Generally, nutrient cycle is closely related to the element distribution in biomass and the population dynamics in ecosystems. Carbon, nitrogen and phosphorus in Japanese anchovy (Engraulis japonicus) of different body lengths from the Huanghai Sea (Yellow Sea) were determined to better understand their variability and reasons during its life history. The mean content was 45.12%, 10.12% and 2.02% for C, N and P, respectively. Significant differences in C, N and P elemental composition were found among different sizes, which could be explained by varying proportions of storage compounds in whole body fish, and varying degrees of ossification. Considering abundant resources in Japanese anchovy, it was an important P-pool in the cycle of P. Moreover, the excreted N/P ratio was significantly different in fishes of different sizes, especially at high gross growth efficiency. In the past two decades, overexploitation tended to cause smaller body length in the community structure of anchovy, which presumably changed the nutrient cycle in food webs of the Huanghai Sea. Exptrapolation of the results indicates that Japanese anchovy may be important for conveying nutrient in the Huanghai Sea.     

表层水温结构变化对东南太平洋秘鲁鳀渔场的影响

秘鲁鳀（Engraulis ringens）是栖息于东南太平洋沿岸的小型中上层鱼类,掌握其渔场变化及其与海洋环境因子的关系有利于企业把握该渔业的生产情况。研究结合2005-2014年渔汛期间秘鲁各港口出港的船数及其所获得的秘鲁鳀渔获量和表层水温数据（海表面温度,sea surface temperature,SST）,以单位捕捞努力量的渔获量（catch per unit effort,CPUE）和捕捞努力量（Effort）数据构建的渔场指数（fishing ground index,FGI）为渔场指标,对秘鲁鳀的渔场类型进行分类,探讨水温结构变化对秘鲁鳀渔场的影响。方差分析表明:渔场指数在不同渔汛阶段（渔汛前期、中期和末期,P<0.01）和不同捕捞区域（北部、中部和南部,P<0.01）都有着极显著的差异。以表层水温大于20℃的海水是否入侵到近岸为标志,可以将秘鲁鳀渔场分成两种类型:大于20℃的海水没有入侵到沿岸（A型渔场）和入侵到沿岸（B型渔场）。研究表明,在渔汛前期和中期阶段,A型渔场的出现有利于渔场的形成,其中渔汛前期沿岸19℃或20℃等温线的出现以及渔汛中期沿岸18℃或19℃等温线的出现可以作为中心渔场形成的指标。     

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.     

Bionomy of the Amphipods in the Evros Delta (North Aegean Sea)

Abstract. The composition and distribution of the amphipod fauna in the area of Evros Delta (North Aegean Sea) have been investigated by means of quantitative sampling. On the basis of similarity in distribution among stations, 3 groups of species have been distinguished. The dependence of the distribution of these 3 groups, as well as of each species separately, on certain environmental factors is discussed.     

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.     

Comparison between amphipod assemblages associated with Caulerpa racemosa var. cylindracea and those of other Mediterranean habitats on soft substrate

The spread of the invasive alga Caulerpa racemosa var. cylindracea in shallow-water habitats can present different faunal assemblage composition. We compared the amphipod assemblages associated with C. racemosa and natural habitats found on shallow-water Mediterranean soft substrata. Four vegetated habitats were compared: C. racemosa, Caulerpa prolifera, Cymodocea nodosa and Posidonia oceanica with unvegetated substrata. Samples were collected during two sampling periods (September 2004 and March 2005). A total of 63 amphipod species were recorded. The results showed that the vegetated habitats sampled, including C. racemosa stands, supported a higher abundance and species richness of amphipods. Furthermore, the assemblage structure differed between the different habitats, while the abundance of some species was significantly different, depending on habitat. For example, Microdeutopus obtusatus was favoured by C. racemosa habitat; Ampelisca diadema was associated with C. prolifera beds; and Hyale schmidti was more abundant in P. oceanica meadows. Habitat invasion by C. racemosa can exert an important influence on biotic assemblages, modifying habitat structure and associated fauna.