Allozymic and morphometric evidence for two stocks of the European anchovy Engraulis encrasicolus in Adriatic waters

Allozyme electrophoresis and morphometric analyses were used to investigate the stock structure of European anchovies (Engraulis encrasicolus L.) captured by Italian vessels in the Adriatic Sea. Twenty four putative enzyme loci were studied, all of which exhibited genotypic proportions in accordance with Hardy-Weinberg predictions. Two loci, IDHP-2 ^* (isocitrate dehydrogenase, 1.1.1.42) and G3PDH-2 ^* (glycerol-3-phosphate dehydrogenase, 1.1.1.8), showed significant allele-frequency differences among samples. IDHP-2 ^* 100 frequencies ranged between 0.35 and 0.88, with lower frequencies recorded in northern waters and areas close to the Italian coastline, while G3PDH-2 ^* 100 frequencies followed a similar, if less distinct, pattern. For both loci, the trends observed were spatially stable over a 2 yr sampling period. A putative stock-boundary was superimposed onto a map of IDHP-2 ^* 100 frequencies, and a bootstrapped dendrogram confirmed the genetic separation of the two putative stocks, one located around the north-western Adriatic and the other in the central-southern region. Canonical variate analysis (CVA) of morphometric data collected using a truss network indicated that the two putative stocks were morphologically distinct; 89.6 and 88.3% of north-western and central-southern anchovies, respectively, were correctly assigned by discriminant-function analysis. This variation may be related to the presence of two anchovy colour phenotypes, silver and blue, in the Adriatic Sea. Silver anchovies are characteristic of northern areas, while the larger blue fish are found mainly in the deeper southern waters. Current assessment models for the fishery are based on the concept of a consistent identity between consecutive catches at single ports, which our data reject. We discuss the possibility of partitioning fishing effort based on the proportions of the two stocks landed at individual ports.     

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.     

Analysis of lipid components for determining the condition of anchovy larvae,Engraulis mordax

It has been presumed that intertidal spawning by Limulus polyphemus minimizes the loss of egges to subtidal predators; however, this strategy involves considerable risks. Massive beach strandings of adults accompany seasonal spawning migrations of crabs along Cape May in Delaware Bay, (USA). At least 190000 horseshoe crabs, approximating 10% of the adult population, died from beach stranding along the New Jersey shore of Delaware Bay during the 1986 (May to June) spawning season. Abnormalities of the telson (which is used in righting behavior) were significantly more common among stranded crabs than among individuals actively spawning on the intertidal beach. The number of stranded crabs per day was not correlated with tidal height or environmental variables (wind speed, wave height) which characterized the conditions at spawning. A complex suite of factors, including the size of the available spawning population, tidal and weather conditions, and beach slope, influence the number stranded during the breeding season. Horseshoe crab stranding results in a large loss of gravid females from the population, and may represent a major input of organic matter to intertidal sandy beaches in certain regions of Delaware Bay.     

Oceanographic habitat,growth and mortality of larval anchovy (<Emphasis Type="Italic">Engraulis encrasicolus</Emphasis>) in the northern Aegean Sea (eastern Mediterranean)

Data from two ichthyoplankton surveys carried out during June 1995 and June 1996 were used to study the broad scale distribution patterns of anchovy eggs and larvae over the northern Aegean Sea continental shelf and the regional/inter-annual variability in growth and mortality rates of larvae. Two major spawning grounds were identified. One in the east, located in the area influenced by the Samothraki gyre (SG), in which a large amount of enriched, modified Black Sea water (BSW) is entrapped and one in the west, associated with zooplankton-rich waters in the semi-enclosed Thermaikos gulf close to several river mouths. In the NE Aegean, anticyclonic gyres generated over the continental shelf and fed by the circulating stream of BSW (like the SG) may act as retention areas for larval anchovy. In the west, the high enclosure of the Thermaikos Gulf contributes to reducing offshore dispersal. Major changes were observed in egg and larval abundance as well as larval mortality between June 1995 and June 1996 in both the western and eastern part of the continental shelf. Mean abundance of eggs and early larvae was >5 times higher in 1996 than in 1995, when waters were significantly cooler, fresher and richer in mesozooplankton. Larval survival decreased from 79 to 69% day⁻¹ in the east and from 89 to 74% day⁻¹ in the west between 1995 and 1996. Hence increased egg production was coupled with higher larval mortality during June 1996. Furthermore, a highly significant positive relationship between larval mortality (Z) and mean egg abundance (A) emerged (Z = −0154 + 0.205 log[A], r ² = 0.96, n = 7) when data from this study and a similar study in the NW Mediterranean were regressed. Mean growth rate of anchovy larvae in the study area (∼0.5 mm day⁻¹) did not differ significantly between areas/years. A marked ontogenetic change was observed in the otolith size/recent otolith growth-on-fish size relationships, which exhibited significant inflection points at ∼6 mm formalin preserved length. This change seems to coincide with performance (e.g., catchability) and behavioral changes (e.g., onset of vertical migrations) in European anchovy associated with the development of the caudal fin (the flexion stage).     

An attempt to determine growth and hatching dates of juvenile sardine,Sardina pilchardus,in the Western Mediterranean Sea

Juvenile sardine, Sardina pilchardus Walbaum, 1792, were collected off Mallorca, Balearic Islands, during April–July 1989, and were aged by means of daily otolith increments. Growth rates were in the range of values reported for other clupeoids, although lower than those for larvae of the Mediterranean anchovy Engraulis encrasicolus. Hatching dates were back-calculated from the age in days and the date of capture. The data indicated that hatching was continuous, but revealed a low peak in the last half of November 1988 and a second, higher peak in the first half of March of the following year, in accordance with the general cycle of secondary production in this oligotrophic area.     

Diets of deep-sea brachyuran crabs in the Western Mediterranean Sea

This study examines the feeding habits of Paromola cuvieri (Risso, 1816) and Geryon longipes A. Milne Edwards, 1881, the only two common deep-sea brachyuran crabs inhabiting the bathyal mud assemblages in the Catalan Sea (Western Mediterranean). Samples were obtained by bottom trawls at depths between 360 and 1871 m during 1983 to 1992. Both species had highly diverse diets, but very low feeding activity, as reflected by the high proportion of empty stomachs. Both characteristics may be important factors enabling deepsea crabs to adapt to bathyal zones, where trophic resources are scarce. The most important food items found in P. cuvieri were fish remains (teleost, sharks) and benthic decapods (Monodaeus couchii, Munida tenuimana). Scavenging activity plays an important role in this species. The diet of G. longipes included a broad range of benthic invertebrates. In the upper middle slope, the bivalve Abra longicallus, decapods (Calocaris macandreae and Monodaeus couchii), echinoderms and polychaetes were the dominant prey, with epibenthic peracarids as a secondary resource. On the lower middle slope, the incidence of decapod crustaceans (C. macandreae, Pontiphilus norvegicus) and peracarids in the diet declined. Small macrobenthic prey (glycerids, cumaceans or amphipods) were rare in the diet of both species, in accordance with the large size of the crab specimens studied. The absence of preferred prey items and the lack of food items of an optimum size on the lower slope may contribute to the progressive decline in abundance of P. cuvieri and G. longipes with increasing depth.     

Importance of the Japanese anchovy (Engraulis japonicus) to breeding rhinoceros auklets (Cerorhinca monocerata) on Teuri Island, Sea of Japan

Temporal variation in the diet and chick growth of rhinoceros auklets (Cerorhinca monocerata), on Teuri Island, Sea of Japan, was studied to understand how local marine environmental changes affect the reproduction of this piscivorous seabird. The food delivered by parents to chicks was sampled every 1-2 weeks from late May to July, 1994-1998. Overall, the diet of nestling rhinoceros auklets consisted of (by mass) 61% Japanese anchovy (Engraulis japonicus), 18% Japanese sand lance (Ammodytes personatus), 18% Japan Sea greenling (Pleurogrammus azonus), 2% other fish and 1% squid. Among years, the contribution of anchovy ranged from 16% to 93%. Once anchovy occurred in the diet, it dominated (80% on average) thereafter. Accordingly, when anchovy appeared in the diet early in the chick-rearing season (1994, 1998), the contribution of anchovy overall was large. The first appearance of anchovy in the diet of auklets late in the summer of 1997 was possibly related to negatively anomalous sea-surface temperature. Food loads composed of anchovy (34.0 g) were heavier than those of sand lance (22.5 g) and greenling (28.5 g). The energy density of anchovies also was higher: 6.3 kJ g^-1 wet mass compared to 0+ greenling (4.78 kJ g^-1) and 0+ sand lance (3.78 kJ g^-1). Thus, a high proportion of anchovy in the diet resulted in high food load mass, high daily growth rates of chicks and high fledging success. This study highlighted the importance of the time of arrival of migratory high-lipid prey, which is influenced by local oceanographic conditions, to the reproductive performance of a piscivorous seabird.     

Nutritional condition and growth rate of anchovy larvae (Engraulis mordax) in the California Current: two contrasting years

A large number of northern anchovy larvae, Engraulis mordax, were collected in the Southern California Bight on two CalCOFI cruises during spring 1987, and the amounts of triacylglycerol, cholesterol and polar lipids were measured in each individual larva using the latroscan TLC/FID (thin-layer chromatography/flame ionization detection) system. The 1987 larvae were compared to larvae taken in the Bight during the previous year. Canonical correlation analyses revealed a significant relationship between the larval lipid components and physical/biological data collected at each occupied station during the late 1987 spring cruise, similar to that recorded in 1986. The short-term nutritional condition of the copepod Calanus pacificus was the most important station variable in both cruises. Their triacylglycerol:cholesterol ratio indicated that few of the 1987 larvae had been starving, but their very low contents of length-specific cholesterol and polar lipid indicated that they had been growing very slowly. Otolith readings on defatted larvae confirmed that the 1987 larvae had been growing significantly more slowly than the 1986 larvae. Furthermore, the data suggested that in the 1986 population the faster growing larvae were also those with the highest lipid content. This situation was reversed for the 1987 larvae, when the faster growing larvae in the population seemed to achieve their faster growth only at the expense of a lowered lipid content. The abundance of anchovy larvae was much higher in spring 1987 than in spring 1986 but, despite this, the apparent recruitment (number of Age 0 fish in July of each year) was not higher in 1987.     

Geographical variations in the trophic ecology of Japanese anchovy, Engraulis japonicus, inferred from carbon and nitrogen stable isotope ratios

Carbon and nitrogen stable isotope ratios of Japanese anchovy (Engraulis japonicus) and their stomach contents were examined and compared among various regions around Japan. Geographical variations in the isotope ratios were found between inshore and Pacific offshore regions. While most of the anchovy samples had isotope ratios around −17.6‰ for δ¹³C and 10.0‰ for δ¹⁵N as median values, higher (more enriched) isotope values were found in the anchovy sampled from inshore regions. On the contrary, lower (more depleted) values were found mostly in the anchovy from the Pacific offshore region including the Kuroshio Extension and Kuroshio-Oyashio transition zones. Higher carbon isotope ratios in the inshore regions may reflect a carbon source from benthic primary producers in addition to phytoplankton possibly through the consumption of the larvae of benthic organisms such as bivalves or decapods, which were found in the stomach contents of the inshore anchovy. Variations in the nitrogen isotope ratio may reflect not only differences in the trophic level of prey species, but also variations in the baseline level of food webs. Stable isotope ratios are potentially a useful tool for understanding the stock/population structure and migration of anchovy. The present findings indicate the potential importance of the “inshore–offshore” variations in the biology of Japanese anchovy populations in the northwestern Pacific waters.     

Tissue distribution and neurotoxic effects of domoic acid in a prominent vector species, the northern anchovy Engraulis mordax

The planktivorous northern anchovy is a prominent vector of the phycotoxin domoic acid (DA) to organisms at higher trophic levels, including fish-eating seabirds and mammals. Although there are abundant data reporting DA-induced excitotoxic symptoms in higher vertebrates, to date there has been no reported evidence of neurotoxic effects in lower vertebrate vectors such as fish. To explain this apparent lack of toxicity, it has been suggested that DA may not reach the brain in anchovies and/or that fish are not as sensitive neurologically to DA. In the present study, intracoelomic (IC) injection of DA, at doses ranging from 1 to 14 μg DA g⁻¹ total fish weight, resulted in severe neurotoxic symptoms such as spinning, disorientation, inability to school, and mortality, indicating that anchovies are neurologically susceptible and that DA crosses the blood–brain barrier in fish. An ED₅₀ of 3.2 μg DA g⁻¹ total body weight was determined via IC injection of DA in 83 anchovies. Comparable intraperitoneal injection studies with mice, rats, and monkeys report similar DA-induced neurotoxic symptoms at doses near 3.2 μg DA g⁻¹, suggesting a similar neurologic sensitivity and mechanism of toxicity between anchovies and mammals. DA tissue distribution measurements from freshly collected field-exposed anchovies and orally gavaged anchovies indicate that DA uptake from the gastrointestinal tract does occur. Levels as high as 1,175 μg DA g⁻¹ were measured in anchovy viscera, while muscle and brain tissue DA levels were 3 orders of magnitude lower, indicating low but measurable DA uptake. Further evidence is needed to confirm that uptake is sufficient during field events to induce symptoms in anchovies. Our work provides the first reported evidence of neurotoxic symptoms in fish and suggests that anchovies may be affected by DA during toxic diatom blooms. If sufficient uptake occurs, DA-induced neurotoxic symptoms and mortality may make fish easier prey targets, thereby selecting for the highest toxin levels transferred, as well as providing a possible pathway for the transfer of DA to benthic communities. Received: 19 May 2000 / Accepted: 29 November 2000     

Vertical distribution of eggs and larvae ofEngraulis encrasicolus in stratified waters of the western Mediterranean

The vertical distribution of the eggs and larvae of the European anchovy (Engraulis encrasicolus) in the western Mediterranean Sea in June 1984 and Agust 1985 was analyzed based on multiple plankton tows carried out at varying depths and using nets equipped with opening and closing mechanisms. Hydrographic parameters such as temperature, salinity, and chlorophylla were recorded simultaneously. Maximum abundance of anchovy eggs and larvae always occurred above the thermocline, even when maximum chlorophylla concentrations were located below the thermocline. Larval distribution appeared to be associated with the availability of suitable food organisms. As in other clupeoid species, the anchovy larvae carried out vertical migrations related to the photoperiod.     

Feeding,growth, and survival of Engraulis mordax larvae reared in the laboratory

Methods are described for the successful rearing of northern anchovy larvae (Engraulis mordax Girard) on cultured foods. Larvae were fed successively on the unarmored dinoflagellate Gymnodinium splendens, the veliger of the gastropod Bulla gouldiana, and nauplii of the brine shrimp Artemia salina. Rearing containers ranging in capacity from 4.5 to 510 l were tested; the smaller ones were found to be most useful for laboratory experimentation. Irreversible starvation occurred when E. mordax were denied food for more than 1.5 days after yolk absorption. Growth rates of larval anchovies fed different diets were compared. Larvae fed G. splendens grew for 1 week at the same rate as animals fed wild plankton, but did not maintain this rate. Laboratory survival of E. mordax larvae on a diet of G. splendens alone, did not differ significantly when veligers supplemented the diet. However, when G. splendens and veligers were fed simultaneously to E. mordax larvae, growth rate was greatly improved, although still not matching the growth attained on a diet of wild plankton. Length (L) versus weight (W) analyses were made for all larvae at all diets. The results showed that weight could be calculated most accurately from length by the relationship log W=3.3237 log L-3.8205, regardless of diet.     

Reproduction,larval development and post-larval growth ofNaineris laevigata (Polychaeta,Orbiniidae) in the Mediterranean Sea

To investigate the life-history adaptations of a brackish-water population ofNaineris laevigata Grube, a population from Acquatina lagoon near Lecce (Italy) was studied. Benthos and plankton samples were collected monthly over a period of ca. 2 yr (1989–1990). Observations of gametogenesis indicate that oogenesis is intraovaric, probably with autosynthetic yolk formation. Oogenesis takes about 7 mo, from November to April–May, for both proliferative and vitellogenetic phases. Reproduction is synchronized within the population. Spawning has also been obtained under laboratory conditions where larval development and post-larval growth were studied. Eggs measured 240 µm in diameter and were spawned within a gelatinous mass. The lecithotrophic pelagic stage was very short. Reproductive effort was high, probably because of high food availability and low level of competition. One of the most interesting findings was the probable presence of a resting stage in the life-cycle. The adaptive significance of enclosing the eggs in gelatiuous masses and of reducing the duration of the pelagic stage is discussed.     

Zooplankton population model coupled to a biogeochemical model of the North Western Mediterranean Sea ecosystem

A stage structured population (SSP) model based on Fennel's [Fennel, W., 2001. Modelling copepods with links to circulation models. Journal of Plankton Research, 23, 1217–1232] equations is applied to Centropages typicus (Kröyer), a dominant copepod species of the North Western Mediterranean Sea (NWMS) and a prey of small pelagic fish. The model considers five groups of stages and development rates are represented by a mechanistic formulation depending on individual specific growth in each stage. Individual growth is calculated from the individual energy budget depending on food availability and temperature.     

Larval development of the euphausiid Euphausia pacifica in the Yellow Sea

The complete larval development of Euphausia pacifica in the Yellow Sea is described and the stages are compared with larvae of E. nana. Euphausiid larvae examined in the present study were collected at 30 stations in the Yellow Sea in 1989. During the stages of Calyptopis III to Furcilia II, E. pacifica larvae are smaller than E. nana larvae but they are larger from Furcilia III onward. There are geographical variations in body size of Calyptopis III among the southern California waters, the eastern Japan waters, the Japan Sea, and the Yellow Sea; body size is smallest in the Yellow Sea and Japan Sea, while it is largest in southern California waters and intermediate in eastern Japan waters. Segmentation of antennal endopods was observed in the 56 to 51 furcilia forms in the Yellow Sea population, suggesting that this characteristic is not as reliable for identification of furcilia stages as the pleopods and terminal telson spines. Thus, we propose here that there are six furcilia stages of E. pacifica, instead of the seven previously reported by Boden (1950).     

Distribution of euphausiid assemblages in the Mediterranean Sea

Eleven species of euphausiids from 24 Isaac-Kidd Midwater Trawl (IKMT) night collections taken at stations throughout the Mediterranean Sea were counted. The frequency of occurrence and dominance of individual species and percent similarity faunal analysis of the euphausiid community were used to describe changes in faunal composition between geographical areas and differences in vertical distribution. Although most species were widespread, three distinct patterns of abundance were apparent: Euphausia krohnii, Nematoscelis megalops, Meganyctiphanes norvegica, and Stylocheiron abbreviatum predominated in western basin areas west of the Tyrrhenian Sea; Euphausia hemigibba, Thysanopoda aequalis, and Stylocheiron longicorne predominated in the Tyrrhenian Sea and eastern Mediterranean Sea; Euphausia brevis and Stylocheiron suhmii predominated in the eastern Mediterranean Sea. Percent similarity analysis of data from the IKMT samples and data from Ruud (1936) indicates the Tyrrhenian Sea fauna at the time of the collections was more similar to eastern Mediterranean areas than to most other areas in the western basin, although the degree of similarity was dependent, to some extent, on the depth at which the samples were collected. The composition of euphausiids living above 150 m at night in this area was more similar to eastern basin areas, while the composition of deeper living forms was more similar to those of the rest of the western basin. Comparison of euphausiids collected at three points over a 60 year time-span in the Balearic Sea shows the similarity in composition to be greater within the area over time than between adjacent areas in the western Mediterranean Sea.Contribution No. 2732 from the Woods Hole Oceanographic Institution, Woods Hole, Massachusetts, USA. This research was supported by the Atomic Energy Commission Contract AT (30-1)-3862, ref. NYO-3862-49, National Science Foundation Grant GA 29303 and Office of Naval Research Contract N00014-71-C-0284.     

A plankton study in the eastern Mediterranean Sea

A synoptic study of the phytoplankton and zooplankton distribution in the eastern Mediterranean Sea was carried out in the summer of 1965 during a cruise of R.V. “Pillsbury”. The phytoplankton maximum was normally found at 100 m, below the 1% level of surface light, with a frequent smaller peak at 40 m. In shallower waters, the main peak was at 80 m. The greatest number of phytophagous crustaceans was recorded at, or close to, the phytoplankton maximum. Depth relationships of phytoplankton and functionally phytoplanktonic species are discussed. Zooplankton was collected from depths down to 4,400 m, the deepest cast being in the Rhodes Deep. The most common species and genera of some selected groups of zooplankton were identified and their distribution considered in the light of some previous cruises in part of the area. The presence of Acantharia containing zooxanthellae with chlorophyll at, and below, 4,000 m was recorded for the first time. Potentially photosynthetic Ceratium vultur and C. carriense (Dinophyceae), both species in active division, were also found at these depths, as well as Halosphaera viridis (Prasinophyceae), which was recorded down to a depth of 1,000 m.