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.     

Metabolism of Centropages species in the Mediterranean Sea and the North Atlantic Ocean

Information on the metabolism rates of Centropages typicus and congeneric species (C. hamatus, C. furcatus, C. brachiatus and C. abdominalis) in neritic areas of the Mediterranean Sea, the North Atlantic Ocean and the Pacific Ocean are reported here. Respiration rates and excretion rates are strongly influenced by abiotic (i.e. temperature, salinity) and biotic factors (i.e. food availability and composition). Differences in the response of respiratory rates to temperature of acclimated, acclimatized and adapted individuals are clearly observed among regions of the Mediterranean Sea and the West and East shores of the Atlantic Ocean. Food supply also strongly affects respiration and excretion rates, as well as the size, sex and stage development of the individuals. The co-measurement of these two rates allows confirmation of the omnivory or carnivory oriented feeding habits of these species. The role of this neritic genus in coastal environment is also discussed.     

The biology and ecology of Centropages typicus: An introduction

We briefly introduce the different chapters of this special issue which review the current knowledge and understanding of the biology and ecology of Centropages typicus obtained over the last 30 years. The papers are grouped into two major theme areas: the first five papers review behaviour, feeding, metabolism, growth and development, and reproduction, and the four following papers have a regional focus dealing with temporal and spatial distribution in the Mediterranean Sea and in the North Atlantic shelf areas. The findings reviewed in the two groups of papers converge to explain why C. typicus is such a very successful copepod species in these coastal areas. In the different chapters, similarities and differences with congeneric species (C. velificatus, C. hamatus, C. brachiatus, C. chierchiae, C. furcatus, and C. abdominalis) have been explored where these are known.     

Macroecological study of Centropages typicus in the North Atlantic Ocean

Centropages typicus is a temperate neritic-coastal species of the North Atlantic Oceans, generally found between the latitudes of the Mediterranean and the Norwegian Sea. Therefore, the species experiences a large number of environments and adjusts its life cycle in response to changes in key abiotic parameters such as temperature. Using data from the Continuous Plankton Recorder (CPR) survey, we review the macroecology of C. typicus and factors that influence its spatial distribution, phenology and year-to-year to decadal variability. The ecological preferences are identified and quantified. Mechanisms that allow the species to occur in such different environments are discussed and hypotheses are proposed as to how the species adapts to its environment. We show that temperature and both quantity and quality of phytoplankton are important factors explaining the space and time variability of C. typicus. These results show that C. typicus will not respond only to temperature increase in the region but also to changes in phytoplankton abundance, structure and composition and timing of occurrence. Methods such as a decision tree can help to forecast expected changes in the distribution of this species with hydro-climatic forcing.     

Turnover time of waters in a deep basin in the Aegean Sea

The Western Gulf is a basin in the Saronikos Gulf. Below about 100 m the water is almost stagnant, essentially isothermal (around 14°C) and it has practically uniform salinity. The level and the mechanism of enrichment in nutrients is discussed. The age of the water estimated from nutrient budget calculations is between one and eight years. The replacement time of water is similar to that found in the Santa Barbara basin, but different from those of the Cariaco Trench and the Black Sea.     

Trans-Atlantic responses of Calanus finmarchicus populations to basin-scale forcing associated with the North Atlantic Oscillation

Populations of the copepod species Calanus finmarchicus often dominate the springtime biomass and secondary production of shelf ecosystems throughout the North Atlantic Ocean. Recently, it has been hypothesised that interannual to interdecadal fluctuations observed in such populations are driven primarily by climate-associated changes in ocean circulation. Here, we compare evidence from the North Sea and Gulf of Maine/Western Scotian Shelf (GoM/WSS) linking fluctuations in C. finmarchicus abundance to changes in ocean circulation associated with the North Atlantic Oscillation (NAO). A particularly striking contrast emerges from this Trans-Atlantic comparison: whereas the North Sea C. finmarchicus population exhibits a negative correlation with the NAO index, the GoM/WSS population exhibits a more complex, positive association with the index. The physical processes underlying these contrasting population responses are discussed in the context of regional- to basin-scale circulation changes associated with the NAO.     

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea,English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C. typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas.     

Spatial and temporal distribution patterns of suspended particulate matter and particulate organic carbon in the Saronikos Gulf (eastern Mediterranean, Greece)

In 1998–1999, beam attenuation coefficient (bac) profiles, suspended particulate matter (SPM) and particulate organic carbon (POC) concentrations were assessed during five cruises in the Saronikos Gulf, eastern Mediterranean, Greece. SPM and POC concentrations (0.05–1.84 mg l⁻¹ and 10.2–468.6 μg l⁻¹, respectively) exhibited strong spatial and temporal variations, related to the different environmental characteristics of various sectors of the gulf, including wind regime and biological productivity. The Elefsis and Keratsini bays, as well as the area around Psyttaleia Island, showed the highest POC concentrations. The vertical distribution of POC at stations in the western basin, as well as in the inner and outer Saronikos Gulf is characterised by higher POC concentrations in surface waters, associated with higher biological activity. The wastewater treatment plant effluents discharged south of the Psyttaleia Island are a major source of organic particles which directly influence the intermediate water layers, at least during the stratification period. Assessments of relationships between bac and SPM or POC concentrations revealed a relatively strong correlation between bac and POC. An equation converting bac readings to POC concentration was established which can be applied to historical and/or future bac measurements, independently of season. POC concentrations estimated from calibrated continuous transmissometer readings were used to estimate the standing stock of POC in the Saronikos Gulf, which varied between 6,110×10⁶ and 13,450×10⁶ g C during the period June 1998 to February 1999.     

Long-term changes in the composition and diversity of deep-slope megabenthos and trophic webs off Catalonia (western Mediterranean): Are trends related to climatic oscillations?

For depths ranging between 650 and 1700 m we have compared recent (2007-2008) to older (from 1988 to 1992) data, searching for long-term changes in the distribution, abundance and composition of deep megafauna (fish and decapods) off the central Catalonian coasts (western Mediterranean). Overall, in the depth interval between 600 and 1100 m, we found higher abundance of fish in 1988-1992 than in 2007, a decrease simultaneous with an increase of decapod crustaceans. Older and more recent haul replicates (after 20 years) had similar assemblage composition in the depth range 1300-1700 m, whereas we found significant changes at 1000 m. Diversity of fish was greater in 1988-1992 than in 2007, while diversity of decapod crustaceans increased between the two periods. Thus, there was a reorganization in benthopelagic communities, rather than a loss of biodiversity. This was in agreement with long-term changes described for diversity of (neritic) zooplankton in the western Mediterranean. We found a dominance of plankton-suprabenthos feeders (e.g. fish such as Lepidion lepidion, Hymenocephalus italicus and Alepocephalus rostratus; the decapods Plesionika spp. and Sergestes arcticus) in 1988-1992. In 2007 by contrast, dominance of plankton-suprabenthos feeders decreased, and assemblages were increasingly composed of benthos-feeding decapods (e.g. Aristeus antennatus, Pontophilus norvegicus and some hermit crabs) preying for instance on polychaetes. These results coincided with low/negative North Atlantic Oscillation index (NAO) in 2007 and in the period immediately before (2004-2006) 2007 (increase of benthos feeders), and with high average NAO in 1988-1992 (decrease of benthos feeders, which in turn may enhance abundance of plankton feeders). The benthic decapod Calocaris macandreae and suprabenthos (small crustaceans, mostly peracarids, living on and just beneath the sediment surface) are key prey in food webs off Catalonian margins, acting as links between surface oceanographic processes and abundance of benthopelagic predators. A conceptual model is presented relating changes in climatic conditions with changes in deep-sea, benthic-suprabenthic food webs. Calocari macandreae was more abundant in 2007, after 3 years of negative NAO. Under negative/low NAO, rainfall and river discharges increase in the NW Mediterranean, which may enhance advective food inputs for macrobenthos. This fits well with the higher abundance of benthos feeders in 2007, as well as with the significant deepening of decapod crustacean assemblages in that same period. Suprabenthos, being short-lived and annual species, were more abundant under positive NAO conditions (1988-1992), probably enhancing the abundance or aggregation of suprabenthic/pelagic feeders.     

The feeding ecology of the copepod Centropages typicus (Kröyer)

Here we report on the current knowledge on the feeding ecology of the planktonic copepod Centropages typicus. We describe the foraging process of C. typicus from the detection of prey to their digestion, considering also the effects of several physical variables on the feeding activity of the species. C. typicus is an omnivorous copepod that feeds on a wide spectrum of prey, from small algae (3–4 μm equivalent spherical diameter, ESD) to yolk-sac fish larvae (3.2–3.6 mm length). It uses both suspensivorous and ambush feeding strategies, depending on the characteristics of the prey. In general, C. typicus exhibits selection for large motile prey, such as ciliates or dinoflagellates, both in nature and laboratory, and this selective pattern is enhanced under moderate intensities of turbulence. Daily rations in the field are somewhat lower than those found in the laboratory, which indicates food limitation. This fact, together with the relatively modest capacity of the species to adapt to fluctuations in food availability may explain the geographical distribution of C. typicus, being restricted to near-shelf waters. In general, it does not seem that C. typicus feeding severely impacts planktonic populations. However, the occasional importance of the species in certain ecosystems is also apparent.     

Parity and disparity between two Chama oysters: the reproductive biology of the Indo-Pacific C. pacifica Broderip,invasive to the Mediterranean Sea; and C. savignyi Lamy,indigenous to the Red Sea

The Indo-Pacific oyster Chama pacifica Broderip, 1835 (Mollusca, Bivalvia, Chamidae) is rarely found in the Northern Red Sea reefs of Eilat (Gulf of Aqaba), where it is outnumbered by its indigenous congener, Chama savignyi Lamy, 1921. The influx of Eritrean biota from the Red Sea into the Mediterranean Sea via the Suez Canal has led to the formation of massive Chama oyster beds along the Eastern Mediterranean shore. However, unlike the Northern Red Sea, the Eastern Mediterranean Chama beds are dominated by C. pacifica oysters, whereas C. savignyi is absent from this region. In an attempt to understand this difference in their respective distribution, the reproductive biology of both species was compared. Histological analysis of the male and female gonads, monitored monthly from March 2009 to August 2010 in both regions, revealed a similar reproductive cycle, comprising six stages: onset of gametogenesis, advanced gametogenesis, ripe, ready to spawn, spent and sexual rest. Female gonads demonstrated an additional, seventh stage – restoration, coinciding with inferred spawning of ripe gametes. Both species were found to be dioecious spawners, with a single, annual, temperature-dependent inferred spawning period. Chama pacifica was found to reproduce efficiently in maximal Mediterranean seawater temperatures not experienced by the Northern Red Sea Chama populations. This study demonstrates the high invasive potential of an oyster species despite its rarity in its source region.     

A multi-trial diagnostic tool in fin whale (Balaenoptera physalus) skin biopsies of the Pelagos Sanctuary (Mediterranean Sea) and the Gulf of California (Mexico)

The main objective of this study was to apply a set of sensitive non-lethal biomarkers in skin biopsies of fin whales (Balaenoptera physalus) to evaluate the toxicological status of this mysticete in the Pelagos Sanctuary (Mediterranean Sea) and in the Gulf of California (Sea of Cortez-Mexico). We developed a “multi-trial diagnostic tool” (based on field and in vitro studies), combining molecular biomarkers (western blot of CYP1A1, CYP2B) and gene expression (qRT-PCR of HSP70, ERα, AHR, E2F-1) with the analysis of OCs, PAHs and PBDEs. The study revealed a higher level of toxicological stress in the Mediterranean fin whales.     

Biology and distribution of hyperiids in the Sea of Okhotsk

Long-term studies in the Sea of Okhotsk (1986–2012) demonstrated that hyperiids account for a small portion of zooplankton, making up only from 1.0 to 5.3% by weight. The minimum hyperiid biomass was observed in the coastal zone. The hyperiid biomass in the open water shelf community increased from spring to autumn. In the Sea of Okhotsk, hyperiids are represented by ten species, among which the most abundant in the northern regions (mainly in Shelikhov Gulf) is Themisto libellula, while the most abundant in other regions is T. pacifica. The distribution of T. pacifica in different seasons showed that the maximum biomass was concentrated in the deep-water zone. The range of T. libellula in the Sea of Okhotsk is mostly limited to the Shelikhov Gulf, but in certain years its habitation area can expand. In warm years with low ice coverage, the maximum expansion of T. libellula to the west in the shelf zone of the northern part of the Sea of Okhotsk was observed. Since variability of salinity and temperature in the northern part of the Sea of Okhotsk may be fatal for the cryophilic stenohaline T. libellula species, forecasted changes in thermohaline circulation will make it possible to predict the population dynamics of this important species.     

First record of a Vestimentifera (Polychaeta: Siboglinidae) from chemosynthetic habitats in the western Mediterranean Sea—Biogeographical implications and future exploration

A new population of vestimentiferan tubeworms was discovered during a recent expedition to a mud volcano field in the Alboran Sea, western Mediterranean Sea. Morphological data and mitochondrial cytochrome-c-oxidase subunit 1 (COI) sequences show that the Alboran tubeworm is essentially identical to Lamellibrachia sp. found in the eastern Mediterranean. This is the first record of a vestimentiferan species in the western basin of the Mediterranean, an area with direct connection to the Atlantic via the Strait of Gibraltar and therefore of great importance to the study of distributional patterns and evolution of Mediterranean species. We examine the current hypotheses on the biogeographic distribution of vestimentiferan species in the eastern Atlantic and Mediterranean Sea and conclude that independently of when Lamellibrachia colonized the Mediterranean, neither the present hydrological settings of both Mediterranean Sea and Atlantic Ocean, nor vestimentiferans reproductive biology are impeditive to the presence of the Mediterranean species of Lamellibrachia in the NE Atlantic. The West African and Lusitanian margins are the most likely places to find living populations of this species in the NE Atlantic.     

Development and growth rates of Centropages typicus

This paper presents an updated review of our knowledge about Centropages typicus development and growth based on a synthesis of all data found in the literature on stage durations, development rates, size, weight, and C:N ratios. Developmental and growth characteristics have been recorded during field time series and cruises, as well as in controlled laboratory experiments. A few laboratory studies have contributed to an understanding of the impact of food quantity, food quality and temperature on growth and development.C. typicus has a sigmoidal development pattern when the contribution of the different stages to total development time is considered, CIV and CV having much longer durations than preceding stages. The development times of this species are closely related to temperature with a nearly equiproportional development. In turn, food conditions, both in quantity and quality, have less effect on development than on growth rate, leading to large variations in weight within stage, even larger than those predicted from changes in length and biovolume. Actual measurements of weight and biochemical composition are required to better understand the growth of C. typicus in relation to food composition. As a consequence, our review shows that weight–length relationships obtained in particular environmental conditions may be inadequate for others.The combination of a temperature-dependent development rate and a food-dependent growth rate may explain why shelf regions are a more favourable environment for C. typicus as compared to offshore regions, as they support large females, production of more and better quality eggs, and probably better survival of the offspring.     

Eastern and Western Poor Cod (Trisopterus minutus capelanus) Populations in the Mediterranean Sea: Evidence from Allozyme and Minisatellite Loci

Abstract. Nine allozyme and two minisatellite loci were used to investigate potential genetic differentiation among three samples of Mediterranean poor cod, Trisopterus minutus capelanus, from the Gulf of Lion, the Tuscan Archipelago and the Aegean Sea. Both types of markers showed consistent results, with F_ST values of 0.0262 and 0.0296 (P < 0.0015, after Bonferroni correction for multiple tests) for allozymes and minisatellites, respectively. Allele frequency heterogeneity tests between pairs of samples showed a clear separation between the two western Mediterranean samples (Gulf of Lion, Tuscan Archipelago) and the eastern one (Aegean Sea). The results indicate that at least two reproductively isolated populations of poor cod occur in the Mediterranean.     

Total organic carbon distribution and budget through the Strait of Gibraltar in April 1998

In order to investigate total organic carbon (TOC) exchange through the Strait of Gibraltar, samples were taken along two sections from the western (Gulf of Cádiz) and eastern (Western Alboran Sea) entrances of the Strait and at the middle of the Strait in April 1998. TOC was measured by using a high-temperature catalytic oxidation method. The results referenced here are based on a three-layer model of water mass exchange through the Strait, which includes the Atlantic inflow, Mediterranean outflow and an interface layer in between. All layers were characterised by a decrease of TOC concentrations from the Gulf of Cádiz to the Western Alboran Sea: from 60–79 to 59–66 μM C in the Atlantic inflow and from 40–60 to 38–52 μM C in the Mediterranean waters, respectively. TOC concentrations in the modified North Atlantic Central Water varied from 43 to 55 μM C. Intermediate TOC values were measured in the interface layer (43–60 μM C). TOC concentrations increased from the middle of the Strait towards continents indicating a contribution of organic carbon of photosynthetic origin along Spain and Morocco coasts or TOC accumulation due to upwelling in the northeastern part of the Strait. Our results indicate that the short-term variability caused by the tide greatly impacts the TOC distribution, particularly in the Gulf of Cádiz. The TOC input from the Atlantic Ocean to the Mediterranean Sea through the Strait of Gibraltar varies from 0.9×10⁴ to 1.0×10⁴ mol C s⁻¹ (or 0.28×10¹² to 0.35×10¹² mol C year⁻¹, respectively). This estimate suggests that the TOC inflow and outflow through the Strait of Gibraltar are two and three orders of magnitude higher than reported via the Turkish Straits and Mediterranean River inputs.     

A unified assessment of marine Mediterranean assemblages: a lesson from benthic hydroids

Hydroids are important components of the communities of rocky bottom shallow coastal areas. The hydrozoan fauna of the Mediterranean Sea is probably one of most investigated in the world, with lots of faunistic and biogeographical studies. However, quantitative studies using the same sampling methodology and controlled sampling effort have been restricted to areas in the Western and Central Mediterranean. We compared hydroid assemblages in four areas of the Mediterranean Sea, from the Gulf of Cádiz to the South Adriatic Sea, following the same sampling and quantification methodology. Our analysis showed the dominant Atlantic character of the assemblages of the Gulf of Cádiz and the South Alboran Sea because of the influence of the inflowing Atlantic waters. Conversely, the hydroid assemblages of the North Alboran Sea were more similar to the assemblages in the Ligurian and in the South Adriatic, and with a number of species ranking between the observed in these two Mediterranean zones. Fourteen species were identified as making the most significant contributions to characterizing the Mediterranean hydroid species pool from a quantitative point of view. These taxa include species previously named as typical components of the Mediterranean hydroid fauna as well as those only recently recorded in the region (Eudendrium moulouyensis) and invaders such as Clytia hummelincki.