Reproductive ecology of Japanese anchovy off the Pacific coast of eastern Honshu, Japan

Spawning frequency of offshore migrant populations of Japanese anchovy Engraulis japonicus ranged from 0·34 to 0·90, and was higher than that of inshore Japanese anchovy. In addition, spawning frequency of offshore populations varied in response to the sex ratio. They spawned at water temperatures of 5.0, 8.6 and 12.6° C, indicating that the minimum critical water temperature of offshore Japanese anchovy was considerably lower than that of inshore populations. A significant positive relationship was observed between water temperature and relative batch fecundity of offshore Japanese anchovy. Furthermore, water temperature was negatively related to egg size, suggesting a trade-o. between relative batch fecundity and egg size for offshore fish. At the same water temperature, relative batch fecundity of offshore populations was higher than that of inshore ones. The comparison of spawning frequency and relative batch fecundity suggests that offshore Japanese anchovy represent a reproductive ecology by which they spawn more eggs than inshore Japanese anchovy within a certain time period.     

Genetic differentiation of European anchovy (Engraulis encrasicolus) along the Moroccan coast reveals a phylogeographic break around the 25th parallel North

Seven microsatellite markers were used to investigate the population structure of the offshore ecotype of the European anchovy (Engraulis encrasicolus) by comparing 12 marine samples collected off the Moroccan coast with an inshore sample taken as a reference for the lagoonal ecotype. F-statistics, correspondence analysis and Bayesian assignment all concurred to cluster the European anchovy in this region into three groups: (i) one reference lagoonal sample, (ii) samples north of the 25°N latitude and (iii) samples south of it. Moreover, the Bayesian cluster analysis pointed toward the existence of an admixture between the group north of 25°N and the lagoonal ecotype, while this was not detectable with the group south of 25°N. Differential introgression between the two ecotypes could be one of the plausible explanations for the observed genetic structure and reveals the possible existence of a phylogeographic break around the 25th parallel North. Our study illustrates the fact that, for those species that encompass several incompletely isolated ecotypes, the level of gene flow among them may vary in space and serve as a tool for stock identification. This information may be useful to improve fishery management of this important harvested species along the Moroccan coast.     

Seaweed biogeography of the mid-Atlantic coast of the United States

The northern boundary of the warm temperate region of the mid-Atlantic coast of the United States is set at Cape Hatteras; the southern boundary lies at Cape Canaveral. There is some spillover of cool temperate species south of Cape Hatteras into North Carolina and spillover of warm temperate species south of Cape Canaveral toward Palm Beach. Elements of the warm temperate flora also extend into the northern Gulf of Mexico, but precise limits to the flora cannot be drawn there. Thirty-one species are endemic to the warm temperate flora. The inshore waters of North Carolina include approximately equal numbers of species with northern and southern centres of distribution; the species of the offshore waters have predominantly southern affinities, but also include most of the endemic species. Seasonal changes in the shallow water flora of North Carolina reflect eurythermal cool temperate and tropical elements in winter and summer respectively and a year-round warm temperate element. These groupings have been verified by experimental studies in which light and temperature were varied. The deep water flora is a summer flora dominated by perennial species. The inshore, eurythermal cool temperate and tropical species have a variety of cryptic stages by which they persist throughout the year.Paper presented at the Seaweed Biogeography Workshop of the International Working Group on Seaweed Biogeography, held from 3–7 April 1984 at the Department of Marine Biology, Rijks-universiteit Groningen (The Netherlands). Convenor: C. van den Hoek.     

Evidence for Distinct Coastal and Offshore Communities of Bottlenose Dolphins in the North East Atlantic

Bottlenose dolphin stock structure in the northeast Atlantic remains poorly understood. However, fine scale photo-id data have shown that populations can comprise multiple overlapping social communities. These social communities form structural elements of bottlenose dolphin (Tursiops truncatus) populations, reflecting specific ecological and behavioural adaptations to local habitats. We investigated the social structure of bottlenose dolphins in the waters of northwest Ireland and present evidence for distinct inshore and offshore social communities. Individuals of the inshore community had a coastal distribution restricted to waters within 3 km from shore. These animals exhibited a cohesive, fission-fusion social organisation, with repeated resightings within the research area, within a larger coastal home range. The offshore community comprised one or more distinct groups, found significantly further offshore (>4 km) than the inshore animals. In addition, dorsal fin scarring patterns differed significantly between inshore and offshore communities with individuals of the offshore community having more distinctly marked dorsal fins. Specifically, almost half of the individuals in the offshore community (48%) had characteristic stereotyped damage to the tip of the dorsal fin, rarely recorded in the inshore community (7%). We propose that this characteristic is likely due to interactions with pelagic fisheries. Social segregation and scarring differences found here indicate that the distinct communities are likely to be spatially and behaviourally segregated. Together with recent genetic evidence of distinct offshore and coastal population structures, this provides evidence for bottlenose dolphin inshore/offshore community differentiation in the northeast Atlantic. We recommend that social communities should be considered as fundamental units for the management and conservation of bottlenose dolphins and their habitat specialisations.     

Responses by four seabird species to a fluctuating availability of Cape Anchovy Engraulis capensis off South Africa

In recent years, Cape Anchovy Engraulis capensis has been the most important food for four seabirds breeding in South Africa–African Penguin Spheniscus demersus, Cape Gannet Morus capensis, Cape Cormorant Phalacrocorax capensis and Swift Tern Sterna bergii. Between 1984 and 1992, biomass of spawning anchovy fluctuated between about 0.5 and 1.75 million tons. Abundance of anchovy was significantly related to numbers of chicks fledged by African Penguins, occurrence of anchovy in the diet of Cape Gannets and numbers of Cape Cormorants and Swift Terns that attempted to breed. Numbers of African Penguins and Cape Gannets that attempted breeding probably also were influenced by abundance of anchovy. African Penguins and Cape Cormorants abandoned nests when anchovy were scarce and deferred breeding until anchovy became more plentiful. Survival of immature African Penguins in a period of anchovy scarcity was enhanced by availability of South African Sardine Sardinops sagax as an alternative food. When anchovy abundance was low, Cape Gannets fed on sardine.     

Diel feeding periodicity, daily ration and vertical migration of juvenile Cape hake off the west coast of South Africa

Feeding periodicity, daily ration and vertical migration of juvenile Cape hake Merluccius capensis are investigated from midwater and bottom trawl collections taken during a 42-h period between 29 February and 2 March 1992 at a fixed position off the west coast of South Africa. Feeding of 10-20 cm hake intensified during the night when they ascended into subsurface layers to prey on recruits of anchovy Engraulis capensis. Larger hake remained close to the bottom, were partially cannibalistic and exhibited no diel feeding periodicity. M. capensis appear to migrate vertically and feed asynchronously in midwater, as individuals, and not as a population, returning to the bottom when satiated. Based upon the exponential rate of decline in stomach fullness throughout the day, the evacuation rate by hake <20 cm was estimated as 0.054 h⁻¹; 90% evacuation of anchovy prey required an estimated 43 h. Using the Elliott & Persson and Eggers methods, the daily ration was estimated as 5.51 and 4.15% of wet body weight respectively. The effect of the foraging behaviour of M. capensis on the appropriateness of acoustic sampling for estimates of their abundance is discussed.     

Possible impacts of environmental change on pelagic fish recruitment: modelling anchovy transport by advective processes in the southern Benguela

Climate change may influence pelagic fish by altering advective processes or by changing where fish choose to spawn. Using a simulation model, the effects of altered advection and spatial distribution of spawning by anchovy on recruitment off South Africa were explored. Cape anchovy Engraulis capensis spawn on the region of the Agulhas Bank, south of South Africa. Currents transport eggs and larvae to nursery areas. Transport of eggs and larvae was modelled using a flow field based on averaged Acoustic Doppler Current Profile data, Feasible Scenarious of altered advection were modelled. For modelling purposes, the ocean surrounding South Africa was divided into blocks with dimensions of a quarter-degree latitude by a quarter-degree longitude. Acoustically measured distributions of spawner biomass for the years 1986–92 were used to calculate egg production per block. In the model, batches were released from each of these blocks each day of the spawning season. The modelling study indicates that passive transport of young anchovy may account for a substantial proportion of year-class variability. Model results show that distribution of spawners influences the distribution of young of the year, as well as the number and the location of advective losses across offshore boundaries.     

To migrate, or not to migrate: partial diel horizontal migration of fish in a temperate freshwater reservoir

The diel horizontal migration (DHM) of fish between the inshore and offshore zones of the ?ímov Reservoir (Czech Republic, deep, stratified, meso-eutrophic) was investigated by a combination of horizontal and vertical hydroacoustic surveys at 3-h intervals over 48 h and day/night purse seining in August 2007. An overwhelming majority of fish were aggregated within the epilimnetic layer. Considering only the horizontal surveys, cyclic diel fish movements between inshore and offshore habitats were apparent, while the total fish biomass remained constant between day and night. A higher fish biomass was detected in the offshore zone during daytime by both hydroacoustics and purse seining. In contrast, a higher fish biomass was recorded at night in the inshore zone. Bream Abramis brama, roach Rutilus rutilus, and perch Perca fluviatilis dominated the daytime offshore fish assemblage whereas bleak Alburnus alburnus prevailed at night. Bream and roach decreased in abundance at night while perch completely disappeared from the offshore habitat. The diel differences in size distributions and direct catches suggested the population-wide horizontal offshore migration of bleak and inshore migration of all perch during dusk. On the other hand, partial inshore migration of bream and roach adults was observed during dusk (52 and 80%, respectively). The different proportions of offshore residents among species and size classes suggested that differences in size, and, therefore, predation vulnerability, contributed to the observed migration patterns.     

Intraspecific variation in palatability and defensive chemistry of brown seaweeds: effects on herbivore fitness

When offered a choice between brown seaweeds (Phaeophyta) from shallow inshore populations versus deeper offshore populations along the mid-Atlantic coast of the United States of America, the herbivorous amphipod Ampithoe longimana consistently preferred plants from the inshore populations. This was the case for three species (Dictyota menstrualis, Spatoglossum schroederi, and Sargassum filipendula) collected from each of a single inshore and offshore site, and for one species (D. menstrualis) collected from each of three inshore and three offshore sites. Bioassay-guided fractionation of chemical crude extracts from D. menstrualis suggested that the relative unpalatability of the offshore plants was due to the lipid-soluble secondary metabolites 4beta-hydroxydictyodial A and 18, O-dihydro-4beta-hydroxydictyodial A 18-acetate, along with minor compounds that were not fully identified. The inshore-offshore pattern did not appear to result from induction of defenses due to herbivory by mesograzers, as mesograzer densities were higher on the more palatable inshore plants. Herbivore feeding preferences for inshore versus offshore seaweeds matched the effects of those seaweeds on their fitness. When juvenile amphipods were raised on inshore versus offshore tissues of D. menstrualis, amphipod survivorship, growth, and ovulation were significantly suppressed on the offshore compared to the inshore tissues. Few previous investigations have studied intraspecific variance in seaweed palatability. We extend these by showing that between-population differences in palatability can persist for several years and by demonstrating that this variance is chemically based and has dramatic effects on herbivore fitness.     

Historical and contemporary factors shape the population genetic structure of the broadcast spawning coral, Acropora millepora, on the Great Barrier Reef

Effective management of reef corals requires knowledge of the extent to which populations are open or closed and the scales over which genetic exchange occurs, information which is commonly derived from population genetic data. Such data are sparse for Great Barrier Reef (GBR) corals and other organisms, with the studies that are available being mostly based on a small number of sampling locations spanning only part of the GBR. Using 11 microsatellite loci, we genotyped 947 colonies of the reef-building coral Acropora millepora from 20 sites spanning almost the full length of the GBR (～12° of latitude and ～1550 km). The results show a major divide between the southernmost central to southern offshore populations and all other sampled populations. We interpret this divide as a signature of allopatric divergence in northern and southern refugia during the Pleistocene glaciations, from which the GBR was subsequently recolonized. Superimposed on this pattern is a cross-shelf genetic division, as well as a separation of inshore populations south of the Cape Clifton Front at ～21.5-22°S. Most inshore populations north of this, as well as mid-shelf populations in the northern and far northern GBR, are open, exchanging recruits frequently. In contrast, inshore populations south of the Cape Clifton Front and offshore populations in the central and southern GBR are largely self-seeding, at least within the spatial resolution that was achieved given our sampling intensity. Populations that have been impacted by recent disturbance events causing extensive coral mortality show no evidence of reduced genetic diversity.     

The use of otolith shape to determine stock structure of <Emphasis Type="Italic">Engraulis encrasicolus</Emphasis> along the Tunisian coast

Engraulis encrasicolus is of great economic importance in the Mediterranean. However, little is known about its stock structure. Otolith shape analysis has been successfully used for fish stock identification. In this study, the stock structure of anchovy caught off the open sea and the coastal area of the Gulf of Tunis, lagoon of Bizerte and Lake of Ichkeul were investigated using otolith shape. Otolith shape was determined by Fourier analysis and then compared among specimens sampled from different areas with forward stepwise canonical discriminant analysis. Significant differences in otolith shape between the open sea and inshore anchovy groups were detected. Otolith shape of anchovy collected in the Lake of Ichkeul was distinct from the other groups. This finding suggests a clear discreteness of the open sea and the continental groups. The data highlighted the potential for using otolith shape analysis for anchovy stock identification, as well as the role of oceanographic features in determining stock separation. These findings will have major implications for anchovy fisheries management in Tunisia. By using a precautionary approach and considering the three areas as separate stocks, fisheries management strategies should be adjusted to achieve optimum sustainable production from each stock and to avoid decreases in genetic variety.     

LONG DISTANCE OFFSHORE MOVEMENTS OF BOTTLENOSE DOLPHINS1

Despite recent progress defining the morphological and genetic characteristics of forms of the bottlenose dolphin inhabiting offshore waters, little is known of their behavior or ranging patterns. Reports suggest that an “offshore form” exists between the 200- and 2,000-m isobaths in distinct Gulf of Mexico and western North Atlantic stocks, while one or more coastal forms inhabit the waters inshore. Two opportunities to track rehabilitated adult male bottlenose dolphins with satellite-linked transmitters occurred in 1997. “Rudy” stranded in NW Florida and was released in the Gulf of Mexico off central west Florida. He moved around Florida and northward to off Cape Hatteras, NC, covering 2,050 km in 43 d. “Gulliver” stranded near St. Augustine and was released off Cape Canaveral, FL. He moved 4,200 km in 47 d to a location northeast of the Virgin Islands. Gulliver swam through 5,000-m-deep waters 300 km offshore of the northern Caribbean islands, against the North Equatorial Current. These records expand the range and habitat previously reported for the offshore stock of bottlenose dolphins inhabiting the waters off the southeastern United States, underscore the difficulties of defining pelagic stocks, illustrate the success of rehabilitation efforts, indicate the value of follow-up monitoring of rehabilitated and released cetaceans, and expand our understanding of the long-range movement capabilities of a dolphin species more commonly thought of as a resident in coastal waters.     

ASSESSMENT OF HETEROGENEITY IN SIGHTING PROBABILITIES OF HUMPBACK WHALES WITHIN VIEWING RANGE OF CAPE VIDAL, SOUTH AFRICA

The migrations of humpback whales on the African east coast were monitored between 1988 and 1991 at Cape Vidal, South Africa. Shore-based surveys of the northward migration were undertaken each winter, and a survey of the southward migration was undertaken in 1990, from an approximately 60-m-high vantage point on a headland. Independent-observer surveys were carried out in both 1990 (22 d) and 1991 (51 d) to determine the proportion of the population within the survey area that were being missed by observers using a single mark-release model. Results were stratified into three distance intervals from the shore and three sighting condition intervals; there were constant sighting probabilities from the south tower under different sighting conditions, while those from the north tower increased slightly as sighting conditions improved. Sighting probabilities from both towers were highest in the intermediate distance interval and decreased in both the inshore and offshore regions.     

Comparison of inshore and offshore fish larval assemblages within the wider Yangtze River estuary (China)

It is hypothesized that fish larvae undertake vertical movements in estuarine waters. To test this hypothesis, a sampling period was selected during times when many spring‐spawning fish reach maximum abundance in coastal waters so that spring and summer variations in larval fish assemblages could be determined in the Yangtze River estuary. Six oceanographic surveys were conducted across the salinity gradient of an inshore (freshwater) and offshore area (brackish intersection) during spring (May) and summer (August) between 2010 and 2012. The fish larval community was dominated by species of Engraulidae, Gobiidae, Champsodontidae and Mugilidae. The pre‐flexion and flexion larval stages of euryhaline marine species, which are dependent on estuaries as nursery areas, were common. The brackish and marine larval assemblage was the most abundant with taxa such as Coilia mystus and Engraulis japonicus accounting for more than 57.3% of the total catch. Spatial differences in the taxonomic composition of larval fish assemblages were evident between the inshore and offshore areas. Additionally, the Yangtze River runoff regulatory functions as affected by the Three Gorges Reservoir operational mode (hydrological alternating operations) showed weak influences on fish and habitat environments. Low salinities from high freshwater inflow limited bay anchovy production in the inshore area.     

The application of an age-structured model to the north Aegean anchovy fishery: an evaluation of different management measures

The objective of this paper is the integration of existing biological and fishery knowledge of anchovy into a unified modelling framework in order to advance our understanding of species' population dynamics under different fishing strategies. The model simulates the anchovy biomass by combining an age-specific growth equation and a continuous age-structured population model based on the McKendrick-Von Foerster equation. Model predictions were compared to the biomass estimates and annual catches during the period 2003-2008. The present work provided direct evidence for the significance of the prespawning period as a critical life period for the management of anchovy stock in the Aegean Sea. It was found that the introduction of additional management measures could increase the profits in the long run for the fishery. However, for these to become apparent they will require a minimum of four years. Results also indicated that the reduction of fishing mortality directed at the spawning stock (recruitment overfishing) and the selective harvesting of younger individuals may be a plausible means of increasing stock's total anchovy biomass. Finally, as a criterion of long-term population survival, we have considered the mathematical notation of persistence. The numerical criteria of persistence in the present model indicated that the anchovy population could be considered viable.     

Mitochondrial and nuclear DNA analyses reveal fine scale geographic structure in bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>) in the Gulf of Mexico

There is a need for biological information to support current stock designations of bottlenose dolphins (Tursiops truncatus) in the Gulf of Mexico. The existence of many inshore, resident “communities” raises questions as to the relationship these dolphins may hold with dolphins inhabiting neighboring inshore and coastal areas. In this study, population subdivision was examined among four resident, inshore bottlenose dolphin stocks (Sarasota Bay, FL, Tampa Bay, FL, Charlotte Harbor, FL and Matagorda Bay, TX) and one coastal stock (1–12 km offshore) in the Gulf of Mexico. Evidence of significant population structure among all areas was found on the basis of both mitochondrial DNA (mtDNA) control region sequence data and nine nuclear microsatellite loci. Estimates of relatedness showed no population contained a significantly high number of related individuals, while separate AMOVAs for males and females indicated that both sexes exhibit a significant level of site philopatry. Results presented here provide the first genetic evidence of population subdivision between the coastal Gulf of Mexico and adjacent inshore areas along the central west coast of Florida. Such strong genetic subdivision is surprising given the short geographical distance between many of these areas and the lack of obvious geographic barriers to prevent gene flow. These findings support the current, separate identification of stocks for bottlenose dolphins inhabiting the eastern coastal and inshore areas of the Gulf of Mexico.     

Small‐scale demographic variation in the stoplight parrotfish Sparisoma viride

Age‐based analysis of the stoplight parrotfish Sparisoma viride was used to examine whether observed differences in their abundance and size structure among reefs in a cross‐shelf portion of the upper Florida Keys could be explained by variation in demographic rates. Annual and daily sagittal otolith increments were enumerated for 176 individuals collected from replicates of reefs in two strata, inshore and offshore reefs (2–6 m depth). von Bertalanffy growth functions fitted to size‐at‐age plots for each site were similar between reefs within each stratum (inshore and offshore), but differed between strata. Sparisoma viride on offshore reefs attained greater average standard length (L_S) at age, greater mean asymptotic size and were longer lived than fish from inshore reefs. Fish on inshore reefs attained only half the maximum age observed on offshore reefs (4 v. 8 years, respectively). No terminal phase fish >4 years of age were found on either reef type. Estimates of mortality rates from age‐frequency data of collected fish revealed higher mortality on inshore reefs. Demographic variables obtained in this study were similar to published values for S. viride from Caribbean reefs but differed significantly from published values from reefs at a similar latitude (Bahamas), reflecting high demographic plasticity on both local and regional scales.     

Evidence for a host role in thermotolerance divergence between populations of the mustard hill coral (Porites astreoides) from different reef environments

Studying the mechanisms that enable coral populations to inhabit spatially varying thermal environments can help evaluate how they will respond in time to the effects of global climate change and elucidate the evolutionary forces that enable or constrain adaptation. Inshore reefs in the Florida Keys experience higher temperatures than offshore reefs for prolonged periods during the summer. We conducted a common garden experiment with heat stress as our selective agent to test for local thermal adaptation in corals from inshore and offshore reefs. We show that inshore corals are more tolerant of a 6‐week temperature stress than offshore corals. Compared with inshore corals, offshore corals in the 31 °C treatment showed significantly elevated bleaching levels concomitant with a tendency towards reduced growth. In addition, dinoflagellate symbionts (Symbiodinium sp.) of offshore corals exhibited reduced photosynthetic efficiency. We did not detect differences in the frequencies of major (>5%) haplotypes comprising Symbiodinium communities hosted by inshore and offshore corals, nor did we observe frequency shifts (‘shuffling’) in response to thermal stress. Instead, coral host populations showed significant genetic divergence between inshore and offshore reefs, suggesting that in Porites astreoides, the coral host might play a prominent role in holobiont thermotolerance. Our results demonstrate that coral populations inhabiting reefs <10‐km apart can exhibit substantial differences in their physiological response to thermal stress, which could impact their population dynamics under climate change.     

Foraging areas of breeding blue-footed boobies Sula nebouxii in northern Peru, as determined by direction recorders

The association between the distribution of blue-footed boobies Sula nebouxii (BFB) and marine productivity around their breeding grounds is unclear. In Peru, they breed in a region seasonally influenced by the cold, nutrient rich Humboldt Current, and by warm, nutrient poor oceanic waters. The foraging range of BFB in Peru is unknown, although some evidence suggest that they may feed in warm offshore waters. During the austral winter 2002 and summer 2003, we tracked 26 BFBs breeding on Isla Lobos de Tierra, Peru (located approximately 15 km west of the mainland), during 59 feeding trips using small direction recorders to determine their marine habitat use. BFBs preferentially moved to the east of the island and foraged over the continental shelf in cold water masses where marine productivity and the abundance of the main prey, the Peruvian anchovy Engraulis ringens , are usually higher than in other areas. Overall, the median maximum foraging distance from the colony was 39 km (range 3–109 km), with 90% of all trips located within 30 km from the mainland, and enclosed in an area of 13,113 km². These results reveal that BFBs exploit the productive waters of the Peruvian Coastal Upwelling System, rather than open oceanic waters as previously suggested. BFBs travelled a greater distance, dispersed in a broader area, and visited more offshore waters during the winter 2002, than during the summer 2003, following the typical seasonal horizontal dispersion of the Peruvian anchovy.     

A new direct method of stock assessment of the loliginid squid

Hydroacoustic research conducted on chokka squid (Loligo reynaudi d’Orbigny, 1845), off the east coast of South Africa from 1994–2005, has led to the development of an innovative stock assessment technique, perhaps applicable to all loliginids that migrate inshore to spawn. This technique combines hydroacoustic biomass estimates made on the spawning concentrations inshore, and minimum biomass estimates made both inshore and offshore using demersal surveys employing the swept-area method. The hydroacoustic estimate uses an improved method to obtain target strength measurements, and squid concentrations are individually mapped from a small boat with a towed transducer. This method may be used even during intense fishing operations because of the manoeuvrability of the small boat inside a tight cluster of fishing vessels. Biomasses of the individual concentrations are then summed. The inshore biomass, also includes dispersed, mature squid migrating between concentrations, this is assessed using a concentration stability factor. The biomass of dispersed squid offshore is again calculated using the swept-area method, a well known demersal survey methodology. The biomass of concentrated (spawning) squid offshore is calculated using the same proportions between concentrated and dispersed squid which were found inshore. All four components are then summed to calculate the total biomass. The result obtained is subject to the effect of complex temporal dynamics, as new animals are recruited to the adult pool and those recently assessed migrate to other sectors of the distribution area.