Site-specific effects on productivity of an upper trophic-level marine predator: Bottom-up, top-down, and mismatch effects on reproduction in a colonial seabird

We investigated the relative roles of bottom-up and top-down factors in limiting productivity of an upper trophic level marine predator. Our primary working hypothesis was that the reproductive success of black-legged kittiwakes (Rissa tridactyla) a piscivorous, colonial-nesting seabird, was most limited by the abundance, distribution, and species composition of surface-schooling forage fishes. A secondary working hypothesis was that reproductive loss to kittiwake nest predators was greatest during years of reduced prey availability. We report on a broad-scale, integrated study of kittiwakes and their prey in Prince William Sound, Alaska. Our study spanned five breeding seasons (1995–1999) and focused on three colonies that differed in size (ranging from ca. 220 to ca. 7000 breeding pairs) and proximity to each other (50–135 km apart). Kittiwakes in PWS encountered a variety of aquatic habitats, creating a complex foraging environment for breeding birds. We measured kittiwake reproductive success and foraging activities, while simultaneously measuring the abundance of surface schooling forage fishes throughout the foraging range of breeding kittiwakes. The abundance of primary prey species for kittiwakes (Pacific herring Clupea pallasi, Pacific sand lance Ammodytes hexapterus, and capelin Mallotus villosus) varied both annually and regionally, with no one region consistently having the greatest abundance of prey. Likewise, kittiwake reproductive success varied considerably among colonies and years.We found that bottom-up, top-down, timing mismatch, and colony-specific effects were all important to kittiwake productivity. Although bottom-up effects appeared to be strongest, they were not evident in some cases until other effects, such as geographic location (proximity of colony to prey concentrations) and top-down predation, were considered. Important bottom-up effects on kittiwake reproductive success were not only total prey abundance and distribution, but also species, age composition, and chronology of prey occurrence (match/mismatch of timing with critical brood-rearing periods); these effects varied by colony.Top-down effects of predation on kittiwake nest contents (independent of prey abundance) confounded seabird-forage fish relationships. Ultimately, when confounding factors were minimized, non-linear asymptotic relationships were identified between kittiwakes and their prey, with an asymptotic threshold of fish school surface area density of ca. 5 m²/km², beyond which top-down, physiological, or phylogenetic constraints likely restrict further reproductive output. The integrated approach of our investigations provided a more thorough understanding of the mechanisms underlying predator–prey relationships in the complex marine environment. However, such mechanistic theories can only be tested and refined through long-term research and monitoring of much greater duration than the 5-year study reported herein.     

Regular habitat switch as an important feeding strategy of an opportunistic seabird species at the interface between land and sea

During deteriorated prey availability, purely pelagic, specialised seabird species have to alter their feeding strategy by extending foraging radii and/or time spent at sea or reducing feeding intervals of chicks. In contrast, more generalised species such as the opportunistic black-headed gull (Larus ridibundus) breeding at the German North Sea coast, can be assumed to react on prey shortages by switching foraging habitats. The coastal zone of the German North Sea provides a rich habitat mosaic consisting of the offshore zone, tidal flats and terrestrial habitats. Thus, we expected distinct temporal and spatial patterns of habitat switch in accordance with prey availability and environmental constraints. We carried out ship-based and aerial surveys as well as dietary analyses and observations on flight activity. We found a significant switch from terrestrial to marine feeding sites both on a daily basis (related to tidal cycle) and over the whole breeding season. Most likely, the latter switch is the result of lower prey availability in the terrestrial habitats and an increasing quality (in terms of prey abundance and energy intake) of the marine area. While there was only moderate variability in habitat use among different years, we revealed significant differences in the diet of birds from different colonies. The high dietary plasticity and flexible feeding strategy, switching between terrestrial and marine prey is certainly of major importance for the success of an opportunistic avian top predator in a complex coastal zone. It is suggested that – compared to situations elsewhere – the number of breeding pairs of black-headed gulls in the German North Sea coast are still stable due to the switch of foraging habitats performed by individuals in this region.     

Ocean control of the breeding regime of the sooty tern in the southwest Indian Ocean

Food availability, which is often seasonal, is regarded as a key factor in the breeding success of seabirds. In oceanic tropical areas, the resources are mostly patchy and ephemeral at the surface, and the seasonality is less marked than at higher latitudes. Such a situation influences greatly the breeding strategies of the oceanic seabird species. We conducted a comparative study of the breeding phenology of the sooty tern (Sterna fuscata) in relation to the local and regional oceanographic conditions around the four major colonies (Europa, Juan de Nova, Lys and Bird Islands) of the southwest Indian Ocean. Over the 1997–2003 period, around all the studied locations, the sea-surface temperature (SST) and the chlorophyll concentration in the Mozambique Channel and the Seychelles area showed clear seasonal differences related to the southern climate and the monsoon phenomena. The breeding activity is synchronized at each studied colony, but the timings are very different. Seasonal reproduction occurs in austral winter at Europa and Bird Island and in austral summer at Juan de Nova; at Lys Island the reproduction is non-seasonal. For the seasonal colonies, there is a large monthly change in SST just before the beginning of reproduction, which is a proxy indicating the annual phytoplankton bloom. This variation is accompanied by the development of oceanic features such as fronts that favour aggregation of prey, and may also play an important role in the presence of schools of surface tuna, which are very important for the foraging success of sooty terns. Conversely, around Lys Island the seasonal variations of the marine environment do not lead to pronounced development of oceanic structures, and consequently, the longer-lasting phytoplankton bloom could explain the non-seasonal breeding regime there. Further studies will help discern the advantages and disadvantages of seasonal and non-seasonal reproduction regime in response to unpredictable fluctuations of the marine environment.     

Monitoring of seabirds in the Benguela upwelling system: can seabirds be used as indicators and predictors of change in the marine environment?

Historical and contemporary data on Benguela seabirds assumed to be responsive to changes in food availability are reviewed as potential indicators and predictors of change in the ecosystem. Cyclic variations in the annual yields of seabird products evident at decadal long time-scales may reflect fluctuations in prey populations at similar periods. However, no such relationship has been demonstrated and consequently the value of such annual yields as indicators is not proved. More recently, interannual variation in diets, foraging effort, breeding success and chick growth of two of the three most abundant species of seabird in the ecosystem have been examined in relation to changes in prey abundance. Although the relative abundance of important forage species in the diets of both species of seabird have reflected regional changes in the absolute abundance of these prey, changes in the values of other parameters thought to reflect food availability were not consistently correlated between and within species. This indicates poor potential for their use as indicators of change at least over the range of abundances experienced in this study. Under present management protocol, data from seabirds do not nor are likely to provide abundance indices that may be used directly in stock assessment of commercially important species. To date, insights into distribution and migration of pelagic fish provided by seabird-derived data probably represent the most valuable contribution of birds to fisheries scientists studying the Benguela system.     

The distribution of pelagic seabirds in relation to the oceanic environment of Gough Island

The distribution and the abundance of flying seabirds are described in relation to the marine environment near Gough Island in the south Atlantic Ocean. Results from a combined seabird, hydrographic and hydro-acoustic survey are assessed in order to highlight possible biotic-abiotic associations. In accordance with recently published work, the macroscale abundance and species composition of avifauna are explained in terms of Gough Island's position between two major biogeographic zones and by the island's utilization as a breeding site for pelagic seabirds. On the mesoscale, seabird abundance was greatest in areas of hydrographical variability, where results from acoustic transects and net hauls suggested concentration of potential food items (i.e. zooplankton and micronekton).     

Community structure across a large-scale ocean productivity gradient: Marine bird assemblages of the Southern Indian Ocean

Our objective was to understand how marine birds respond to oceanographic variability across the Southern Indian Ocean using data collected during an 16-day cruise (4–21 January 2003). We quantified concurrent water mass distributions, ocean productivity patterns, and seabird distributions across a heterogeneous pelagic ecosystem from subtropical to sub-Antarctic waters. We surveyed 5155 km and sighted 15,606 birds from 51 species, and used these data to investigate how seabirds respond to spatial variability in the structure and productivity of the ocean. We addressed two spatial scales: the structure of seabird communities across macro-mega scale (1000 s km) biogeographic domains, and their coarse-scale (10 s km) aggregation at hydrographic and bathymetric gradients. Both seabird density and species composition changed with latitudinal and onshore–offshore gradients in depth, water temperature, and chlorophyll-a concentration. The average seabird density increased across the subtropical convergence (STC) from 2.4 birds km⁻² in subtropical waters to 23.8 birds km⁻² in sub-Antarctic waters. The composition of the avifauna also differed across biogeographic domains. Prions (Pachyptila spp.) accounted for 57% of all sub-Antarctic birds, wedge-tailed shearwaters (Puffinus pacificus) accounted for 46% of all subtropical birds, and Indian Ocean yellow-nosed albatross (Thallasarche carteri) accounted for 32% of all birds in the STC. While surface feeders were the most abundant foraging guild across the study area, divers were disproportionately more numerous in the sub-Antarctic domain, and plungers were disproportionately more abundant in subtropical waters. Seabird densities were also higher within shallow shelf-slope regions, especially in sub-Antarctic waters, where large numbers of breeding seabirds concentrated. However, we did not find elevated seabird densities along the STC, suggesting that this broad frontal region is not a site of enhanced aggregation.     

Interannual variations of the occurrence of epipelagic fish in the diets of the seabirds breeding on Teuri Island, northern Hokkaido, Japan

The diets of breeding seabirds can be a good monitor of marine environmental changes. From 1984 to 2001 we monitored the diets of black-tailed gulls (Larus crassirostris) (“surface foragers”), rhinoceros auklets (Cerorhinca monocerata) (“epipelagic divers”), and Japanese cormorants (Phalacrocorax filamentotus) (“bottom divers”) that breed on Teuri Island at the northern boundary of the Tsushima Warm current in the Sea of Japan/East Sea. Between 1984 and 1987, both the gulls and the auklets foraged on the sardine (Sardinops melanostictus), but after 1992, they switched to the anchovy (Engraulis japonica). This change might reflect the collapse of the sardine stock in the late 1980s. In the 1990s, the year-to-year variations of the percentage of anchovy in the diets of the three seabird species showed similar trends: High in 1994 and 1998–2001; and low in 1992–1993 and 1995–1997. The estimated stock size of the anchovy population in the Tsushima Current area was positively correlated with the percentage of mass of anchovy in the seabirds’ diets. Thus, the short-term annual changes of the total anchovy availability, which might reflect SST or the volume transport of Tsushima Current, possibly affected the seabirds diets on this island.     

The dynamics of grey seals (Halichoerus grypus) off Amrum in the south-eastern North Sea — evidence of an open population

Grey seals (Halichoerus grypus) have been extinct in the south-eastern North Sea for centuries. Only in recent decades have they re-established a few colonies, the first of which was located on a group of sand bars off the German island of Amrum. Based on frequent counts from aircraft and boats in 1976–2000, and a photo-identification program in 1997, we investigated seasonal dynamics, trends in abundance, pup production, and pup survival in this population. Numbers of hauled-out seals were highest in spring, with a peak of 70–100 in late March or early April, and usually lower than 40 from late May to late January, including the winter breeding season. Both spring and summer counts indicated an average increase of 4–5% per year. Annual pup production was 3–10 in 1988–1995 and 9–13 in 1996–1999, suggesting a breeding stock of only about half the minimum total population size, i.e. 137 individually recognised seals in 1997. The estimated survival of pups until weaning seemed high enough to support autochthonous increase, and hence suggested a self-sustaining breeding stock at least until the late 1990s. Pup survival declined, however, from 95–100% in 1988–1991 to 60–70% in 1992–1999, probably due to erosion of the sand bars. From (i) the relative excess of hauled-out seals in spring, as compared to the phenology of stocks in the western North Sea, (ii) turnover and net loss of individuals during the spring and summer of 1997, as indicated by photo-ID records, and (iii) the low apparent per capita birth rate, we suggest that grey seals off Amrum split into a resident breeding stock and a similar or higher number of seals immigrating outside the breeding season. The latter may originate from larger colonies in the southern and western North Sea, and return there for breeding. In the future, grey seals currently breeding off Amrum may eventually leave the area because of deteriorating conditions. However, a decline of numbers occurring in spring and summer would not necessarily be expected, because seasonal influx from increasing stocks elsewhere in the North Sea is likely to continue.     

Application of a Conceptual Ecological Model to Predict the Effects of Sand Mining around Chilsan Island Group in the West Coast of Korea

Chilsan Island Group, located on the west coast of South Korea, has been recognized as a critical breeding and nursery ground for endangered seabirds because food is abundant and human activity is low. Chilsan Island Group has been under protection as a Natural Monument (Article no. 389) since 1997, but attempts were made in 2004 to exploit the coastal sands around the islands. A conceptual ecological model (CEM) was employed with energy flow diagram (EFD) to predict the effects of sand mining on the islands’ ecosystem. The results showed that sand mining activities caused long-term damage to benthic ecosystems and threatened seabird communities by reducing fish populations throughout the food web. The changes in energy flow in the ecosystem due to sand mining operations predict that reduction in secondary production of benthic animal communities might reduce energy transfer to humans and seabirds by 30%. Different ranges of uncertainty provided by the Monte Carlo simulation lead to the conclusion that sand mining could bring about undesirable consequences. One concern is that such negative consequences would have a much more severe impact on seabirds than humans because seabirds cannot compete effectively against humans for the limited food sources. In this study, the complex interrelationships between ecosystem members were simplified and a CEM was established to clearly identify which elements would be affected. The CEM with EFD was found to be helpful in promoting communication between stakeholders, and it is expected to be widely used as a decisionmaking tool by public officials.     

Protection of marine birds and turtles at St Brandon’s Rock,Indian Ocean,requires conservation of the entire atoll

A survey of seabirds and turtles at St Brandon’s Rock, 400 km north of Mauritius, was undertaken in 2010. We estimated that 1 084 191 seabirds comprising seven breeding species and excluding non-breeders were present at the archipelago and we counted 279 turtle tracks and nesting pits of green turtles Chelonia mydas. Hawksbill turtles Eretmochelys imbricata were also present. Analyses of 30 different islets that make up the atoll showed that the seabird species mostly partitioned their use of islets based on islet size, with four species preferring larger islets and two species preferring smaller islets. Alien species introduced historically are still present and other threats, such as shipwrecks, remain. We propose conservation and other measures that should adequately protect the birds, turtles and coral reef by treating the atoll as a system.     

High Arctic sea ice conditions influence marine birds wintering in Low Arctic regions

Ocean climate change is having profound biological effects in polar regions. Such change can also have far-reaching downstream effects in sub-polar regions. This study documents an environmental relationship between High Arctic sea ice changes and mortality events of marine birds in Low Arctic coastal regions. During April 2007 and March 2009, hundreds of beached seabird carcasses and moribund seabirds were found along the east and northeast coasts of Newfoundland, Canada. These seabird “wrecks” (i.e. dead birds on beaches) coincided with a period of strong, persistent onshore winds and heavily-accumulated sea ice that blocked bays and trapped seabirds near beaches. Ninety-two percent of wreck seabirds were Thick-billed Murres (Uria lomvia). Body condition and demographic patterns of wreck murres were compared to Thick-billed Murres shot in the Newfoundland murre hunt. Average body and pectoral masses of wreck carcasses were 34% and 40% lighter (respectively) than shot murres, indicating that wreck birds had starved. The acute nature of each wreck suggested that starvation and associated hypothermia occurred within 2–3 days. In 2007, first-winter murres (77%) dominated the wreck. In 2009, there were more adults (78%), mostly females (66%). These results suggest that spatial and temporal segregation in ages and sexes can play a role in differential survival when stochastic weather conditions affect discrete areas where these groups aggregate. In wreck years, southward movement of Arctic sea ice to Low Arctic latitudes was later and blocked bays longer than in most other years. These inshore conditions corresponded with recent climate-driven changes in High Arctic ice break-up and ice extent; coupled with local weather conditions, these ice conditions appeared to be the key environmental features that precipitated the ice-associated seabird wrecks in the Low Arctic region.     

Seabirds in the diet of Cape fur seals Arctocephalus pusillus pusillus at three mainland breeding colonies in Namibia

Predation by Cape fur seals Arctocephalus pusillus pusillus on seabirds has previously been attributed to a few individuals, mainly males. Scat samples were collected at three mainland breeding colonies of seals in Namibia to determine the extent of seabird predation. Mainly females were sampled. No scats from the van Reenen Bay and Atlas-Wolf Bay colonies, and only two from Cape Cross, contained feather remains, giving an overall frequency of occurrence of feathers in scats of 0.1%. The feathers found were of African penguin Spheniscus demersus and a species of cormorant (Phalacrocorax sp.).     

No microzooplankton grazing during a Mediopyxis helysia dominated diatom bloom

The new diatom species Mediopyxis helysia was described to science from clones found in 2003 in the North Sea, northern Wadden Sea, and the Gulf of Maine. Seven years after its first occurrence, we observed Mediopyxis to contribute up to almost 50% of the biovolume of the diatoms during a diatom spring bloom in the western Wadden Sea. Grazing experiments based on the dilution technique could not detect any microzooplankton grazing impact on the bloom community. Mediopyxis is now also well established in the western Wadden Sea and does have the potential to become a dominant species. The reasons for its success remain largely unresolved but avoidance of being grazed might be one factor. Future research on this new species is needed to understand the success and forecast the ecological footprint of this large diatom species arriving in the western European Seas.     

Patterns and trends in seabird bycatch in the pelagic longline fishery off South Africa

Both foreign and domestic pelagic longline fishing vessels operate in South Africa’s Exclusive Economic Zone and adjacent international waters where they kill hundreds of seabirds each year as bycatch. To update assessments of the impact of the pelagic longline fishery on seabirds off South Africa, information on necropsied seabirds and national fisheries observer bycatch records were summarised for 2006–2013. Foreign-flagged (Asian) vessels had 100% observer coverage throughout the study period, whereas only 6% of the fishing effort by South African-flagged vessels was observed (with no coverage in 2011–2013). Vessels with observers caught seabirds at a rate of 0.132 birds per 1 000 hooks, resulting in an estimated mortality of 2 851 individuals (356 per year) comprising 14 species. Extrapolation of the observed fishing sets to the unobserved fishing sets by the South African domestic longline fleet suggested that approximately 750 additional birds were likely killed during the study period, therefore a combined 450 birds were killed per year. White-chinned petrel Procellaria aequinoctialis was the most frequently killed species (66%), followed by ‘shy-type’ albatrosses Thalassarche cauta/steadi (21%), black-browed albatross T. melanophris (7%), Indian yellow-nosed albatross T. carteri (3%), and Cape gannet Morus capensis (2%). The seabird bycatch rates were lower than in 1998–2005. Nationality of the vessel, time of line-setting, moon phase, year, season, fishing area, and seabird bycatch mitigation measures all influenced seabird mortality. Concurrent with 100% observer coverage, significant reductions in the seabird bycatch rate occurred in the Asian fleet in the latter years of the study, and these rates now approximate the national target (0.05 birds per 1 000 hooks). However, seabird bycatch rates remained high in the South African fleet, where no observers were deployed during 2011–2013, highlighting the need for independent observer programmes in fisheries—a matter of global interest. Suggestions are made as to how seabird bycatch by pelagic longline fisheries off South Africa may be further reduced.     

A hard-knock life: the foraging ecology of Cape cormorants amidst shifting prey resources and industrial fishing pressure

Once one of the most numerous seabirds of the Benguela upwelling system, the population of Cape cormorants Phalacrocorax capensis has decreased by 60% in the past three decades and the species is listed as Near Threatened. Declines in prey availability and/or abundance brought about by recent changes in the distribution of pelagic fish stocks and industrial purse-seine fishing are hypothesised to be a key driver of seabird population decreases in the southern Benguela. We investigated the foraging behaviour of breeding Cape cormorants by deploying GPS and temperature–depth recorders on 24 breeding adults from three islands off the coast of South Africa, two of them to the north of Cape Point and a third farther south on the western Agulhas Bank. This provided the first measures of foraging dispersal by a cormorant in the Benguela system, and enabled a comparison of foraging behaviour between birds from these islands. Foraging trips of Cape cormorants lasted between 17 min and >7 h, at a maximum distance of between 2 and 58 km away from their colony. Foraging effort was significantly greater for birds from farther north off the West Coast in terms of trip duration, distance travelled, number of dives and time spent flying compared to those from the southernmost island (Dyer), which is probably a response to low prey availability in the north. Coastal reserves that exclude pelagic fishing from inshore feeding grounds around Cape cormorant breeding colonies may result in increased local prey availability, which would benefit Cape cormorant populations.     

Top-down and bottom-up factors affecting seabird population trends in the California current system (1985–2006)

To characterize the environmental factors affecting seabird population trends in the central portion of the California current system (CCS), we analyzed standardized vessel-based surveys collected during the late spring (May–June) upwelling season over 22 yr (1985–2006). We tested the working hypothesis that population trends are related to species-specific foraging ecology, and predicted that temporal variation in population size should be most extreme in diving species with higher energy expenditure during foraging. We related variation in individual species abundance (number km^?2) to seasonally lagged (late winter, early spring, late spring) and concurrent ocean conditions, and to long-term trends (using a proxy variable: year) during a multi-decadal period of major fluctuations in the El Niño-Southern oscillation (ENSO) and the Pacific decadal oscillation (PDO). We considered both remote (Multivariate ENSO Index, PDO) and local (coastal upwelling indices and sea-surface temperature) environmental variables as proxies for ocean productivity and prey availability. We also related seabird trends to those of potentially major trophic competitors, humpback (Megaptera novaeangliae) and blue (Balaenoptera musculus) whales, which increased in number 4–5-fold midway during our study. Cyclical oscillations in seabird abundance were apparent in the black-footed albatross (Phoebastria nigripes), and decreasing trends were documented for ashy storm-petrel (Oceanodroma homochroa), pigeon guillemot (Cepphus columbus), rhinoceros auklet (Cerorhinca monocerata), Cassin’s auklet (Ptychoramphus aleuticus), and western gull (Larus occidentalis); the sooty shearwater (Puffinus griseus), exhibited a marked decline before signs of recovery at the end of the study period. The abundance of nine other focal species varied with ocean conditions, but without decadal or long-term trends. Six of these species have the largest global populations in the CCS, and four are highly energetic, diving foragers. Furthermore, three of the diving species trends were negatively correlated with the abundance of humpback whales in the study area, a direct competitor for the same prey. Therefore, on the basis of literature reviewed, we hypothesize that the seabirds were affected by the decreasing carrying capacity of the CCS, over-exploitation of some prey stocks and interference competition from the previously exploited, but now increasing, baleen whale populations. Overall, our study highlights the complexity of the ecological factors driving seabird population trends in the highly variable and rapidly changing CCS ecosystem.     

Variation in the occurrence of marine birds at fronts in the Bering Sea

Topographically-induced flow gradients can concentrate seabird prey in several different ways, and are potentially important to marine birds in shelf ecosystems such as the south-eastern Bering Sea. We tested the hypothesis that attendance by seabirds was greater at fronts maintained by strong surface flow gradients than at fronts maintained by weak or intermittent surface flow gradients. An analysis of 62 crossings of flow gradients identified from temperature and salinity gradients showed that local increase in attendance was greater in areas of strong surface flow gradient than in an area of weak surface flow gradients. Attendance by marine birds depended on the strength of a flow gradient rather than on the presence or absence of a front.     

How well is the EU protecting its seabirds? Progress in implementing the Birds Directive at sea

The European Commission has repeatedly requested its Member States to fully implement one of the world's oldest nature legislation laws, the Birds Directive, by designating Special Protection Areas (SPAs) in the marine environment. The current work analyses the spatial coverage offered by the SPA network to the 82 species of seabirds that occur in EU waters, and compare it to that of Important Bird and Biodiversity Areas (IBAs), as defined by BirdLife International. Marine SPAs represent 3.9% of the EU's marine area, while the IBA network represents 4.4%. On average, only 0.05% of the EU marine area is protected, and there are enormous regional differences. On average, SPAs and IBAs both cover 16.0% of the breeding distribution range of seabird species, but only1.4% of their non-breeding distribution range. SPAs cover more than 50% of the breeding range of only one seabird family, Procellariidae, and both SPA and IBA networks offer greater coverage to threatened species than to non-threatened ones. The year when Member States were legally obliged (2008 onwards) to declare their marine SPA networks is not significantly related to their network size, and that those species subjected to tracking studies or have well known breeding colonies benefit from higher coverage. Methodological challenges that were once posed to implement the Birds Directive at sea are largely overcome, and political will is the main driver for identifying, designating and managing a complete network of marine SPAs in the EU.     

Accounting for indirect effects and non-commensurate values in ecosystem based fishery management (EBFM)

Ecosystem-based fishery management (EBFM) requires taking account of indirect effects (such as habitat destruction, incidental mortality, and competition between the fishery and marine mammals or birds) and dealing with non-commensurate values (such as yield from the fishery and production of offspring by the birds or mammals competing for the same resource). The perspective of EBFM requires that the rate of fishing mortality is less than the value that provides maximum sustainable yield (MSY), but the question is how far below this level should the fishery operate? For this problem in multiobjective programming, simple method of solution was developed and illustrated with the fishery for sandeels (Ammodytes spp.) in the Shetland Islands. The yield from the fishery at a given fishing mortality F is scaled by MSY (so that this quantity increases as fishing mortality increases from 0 to that giving MSY) and the breeding success of predators (black-legged kittiwakes Rissa tridactyla and Arctic terns Sterna paradisaea) at a given fishing mortality is scaled by that in the absence of fishing. The result is two non-dimensional quantities that can be combined into a single value function, which can then be explored or optimized. It is shown that a reduction of only about 20 percent in yield can nearly double the breeding performance of the more sensitive predator. Extensions of the method are discussed; these include the use of maximum economic yield (MEY) and state dependent life history, as implemented by stochastic dynamic programming.     

Reproduction,hatching success,and early naupliar survival in Centropages typicus

The broadcast spawner, Centropages typicus, is a very successful copepod species in many coastal areas of the Mediterranean Sea and North Atlantic Ocean. This review assembles the large amount of information on the reproduction and early life history of C. typicus that has emerged since the 1970s and has made this species one of the best-studied copepods, similar in that regard to species of Acartia and Calanus. Observations on mating behavior and the female gametogenic and oogenic cycles are presented, together with information on seasonal cycles of egg production rates in Mediterranean and Atlantic populations from various regions. These studies indicate a strong latitudinal gradient, with continuous reproduction and the main spawning season occurring earlier (late winter/spring) in warmer waters such as the Mediterranean Sea, compared to northern areas such as the North Sea and in the Kattegat, where C. typicus actively reproduces mainly in late summer and fall with reproduction ceasing altogether in winter in the German Bight. These observations strongly suggest that temperature is the controlling factor for reproductive activity in this species. Egg development times are also temperature dependent but do not vary with latitude, and there is as yet no conclusive evidence that diapause egg production occurs in C. typicus. Laboratory experiments have shown that food quantity and quality both affect fecundity and offspring fitness, but most of these studies have focused on diatom and dinoflagellate diets and non-algal prey have been strongly underrepresented, despite their importance for this omnivorous copepod. Large fluctuations in hatching success and naupliar survival have been reported in field surveys and have subsequently been related to maternal feeding history and food quality or toxicity in laboratory experiments. We identify future lines of research that will help to explain the interannual variability in breeding intensity and recruitment of C. typicus in the field based on better understanding of the species’ reproductive biology.