Climate change impact on Balearic shearwater through a trophic cascade

A recent study showed that a critically endangered migratory predator species, the Balearic shearwater Puffinus mauretanicus, rapidly expanded northwards in northeast Atlantic waters after the mid-1990s. As a significant positive correlation was found between the long-term changes in the abundance of this seabird and sea temperature around the British Isles, it was hypothesized that the link between the biogeographic shift and temperature occurred through the food web. Here, we test this conjecture and reveal concomitant changes in a regional index of sea temperature, plankton (total calanoid copepod), fish prey (anchovy and sardine) and the Balearic shearwater for the period 1980-2003. All three trophic levels exhibit a significant shift detected between 1994 and 1996. Our findings therefore support the assertion of both a direct and an indirect effect of climate change on the spatial distribution of post-breeding Balearic shearwater through a trophic cascade.     

Migration pathways, behavioural thermoregulation and overwintering grounds of blue sharks in the Northwest Atlantic

The blue shark Prionace glauca is the most abundant large pelagic shark in the Atlantic Ocean. Although recaptures of tagged sharks have shown that the species is highly migratory, migration pathways towards the overwintering grounds remain poorly understood. We used archival satellite pop-up tags to track 23 blue sharks over a mean period of 88 days as they departed the coastal waters of North America in the autumn. Within 1-2 days of entering the Gulf Stream (median date of 21 Oct), all sharks initiated a striking diel vertical migration, taking them from a mean nighttime depth of 74 m to a mean depth of 412 m during the day as they appeared to pursue vertically migrating squid and fish prey. Although functionally blind at depth, calculations suggest that there would be a ~2.5-fold thermoregulatory advantage to swimming and feeding in the markedly cooler deep waters, even if there was any reduced foraging success associated with the extreme depth. Noting that the Gulf Stream current speeds are reduced at depth, we used a detailed circulation model of the North Atlantic to examine the influence of the diving behaviour on the advection experienced by the sharks. However, there was no indication that the shark diving resulted in a significant modification of their net migratory pathway. The relative abundance of deep-diving sharks, swordfish, and sperm whales in the Gulf Stream and adjacent waters suggests that it may serve as a key winter feeding ground for large pelagic predators in the North Atlantic.     

Chaos and stability of age-0 fish assemblages in a temperate deep reservoir: unpredictable success and stable habitat use

Large year-to-year variability in different fish species recruitment has been confirmed by previous studies while diurnal patterns of occupation in two basic reservoir habitats (pelagic and littoral) by different age-0 fish species in late summer are still unclear. Data collected over an 11-year period regarding late-summer age-0 fish assemblages in pelagic and littoral habitats of a reservoir were used to test the recruitment instability and to investigate diurnal habitat use. Trawling was conducted in the pelagic habitat at night while beach seining was conducted in the littoral habitat during day and night. Fluctuations in age-0 fish abundance and species composition were observed with both sampling methods; however, the following spatio-temporal patterns were relatively stable in most investigated years: (1) pelagic species (pikeperch; Sander lucioperca, small perch; Perca fluviatilis, bream; Abramis brama at night), (2) littoral species (large perch, asp; Leuciscus aspius, dace; Leuciscus leuciscus), (3) migratory species likely performing diel horizontal migrations (bleak; Alburnus alburnus), (4) species abundant in the littoral habitat both during day and night and also in pelagic habitat at night (roach; Rutilus rutilus) and (5) species detected in both habitats exclusively at night (ruffe; Gymnocephalus cernuus).     

Spawning Behaviour and Post-Spawning Migration Patterns of Atlantic Bluefin Tuna (Thunnus thynnus) Ascertained from Satellite Archival Tags

Spawning behaviour of Atlantic bluefin tuna (Thunnus thynnus) was investigated using electronic satellite tags deployed in the western Mediterranean spawning ground, around the Balearic Islands (years 2009-2011). All the fish were tagged underwater and released within schools. In general, the fish tagged in the same year/school displayed common migratory trends. Following extended residency around the Balearic Islands, most tagged tuna crossed the Strait of Gibraltar heading for the North Atlantic. Discrepancies between the migratory tracks reconstructed from this and previous electronic tagging studies suggest that the bluefin tuna Mediterranean population may comprise distinct units exhibiting differing migratory behaviours. The diving behaviour varied between oceanic regions throughout the migratory pathways, the shallowest distribution taking place in the spawning ground and the deepest at the Strait of Gibraltar. A unique diving pattern was found on the majority of nights while the fish stayed at the spawning ground; it consisted of frequent and brief oscillatory movements up and down through the mixed layer, resulting in thermal profiles characterized by oscillations about the thermocline. Such a pattern is believed to reflect recent courtship and spawning activity. Reproductive parameters inferred from the analysis of vertical profiles are consistent with those estimated in previous studies based on biological samples.     

Biocomplexity in a highly migratory pelagic marine fish, Atlantic herring

The existence of biologically differentiated populations has been credited with a major role in conferring sustainability and in buffering overall productivity of anadromous fish population complexes where evidence for spatial structure is uncontroversial. Here, we describe evidence of correlated genetic and life history (spawning season linked to spawning location) differentiation in an abundant and highly migratory pelagic fish, Atlantic herring, Clupea harengus, in the North Sea (NS) and adjacent areas. The existence of genetically and phenotypically diverse stocks in this region despite intense seasonal mixing strongly implicates natal homing in this species. Based on information from genetic markers and otolith morphology, we estimate the proportional contribution by NS, Skagerrak (SKG) and Kattegat and western Baltic (WBS) fish to mixed aggregations targeted by the NS fishery. We use these estimates to identify spatial and temporal differences in life history (migratory behaviour) and habitat use among genetically differentiated migratory populations that mix seasonally. Our study suggests the existence of more complex patterns of intraspecific diversity than was previously recognized. Sustainability may be compromised if such complex patterns are reduced through generalized management (e.g. area closures) that overlooks population differences in spatial use throughout the life cycle.     

Climate-driven range expansion of a critically endangered top predator in northeast Atlantic waters

Global climate change is driving rapid distribution shifts in marine ecosystems; these are well established for lower trophic levels, but are harder to quantify for migratory top predators. By analysing a 25-year sightings-based dataset, we found evidence for rapid northwards range expansion of the critically endangered Balearic shearwater Puffinus mauretanicus in northeast Atlantic waters. A 0.6 degrees C sea surface temperature increase in the mid-1990s is interpreted as an underlying controlling factor, while simultaneous northward shifts of plankton and prey fish species suggests a strong bottom-up control. Our results have important conservation implications and provide new evidence for climate-driven regime shift in Atlantic ecosystems.     

Patterns in species richness, size, and latitudinal range of East Atlantic fishes

Examination of latitudinal patterns in species richness, size, and distributional range of East Atlantic fish, based on a compilation of data encompassing the full latitudinal and depth distribution of 1746 East Atlantic fish species, showed that species richness declined towards higher latitudes at a rate of c 1 % of the number of species present, in five-degree bands, for each degree of latitude for both teleosts and elasmobranchs, regardless of habitat However, the latitudinal patterns in maximum fish size and latitudinal range differed between teleosts and elasmobranchs, and changed with habitat No clear evidence was obtained that the latitudinal range occupied increased with latitude, indicating that Rapoport's rule does not apply to E Atlantic fishes Rather, the latitudinal patterns in species richness, size, and distributional range of benthic Atlantic fish were depth-dependent, because species richness, average maximum size, and the average latitudinal range increased with depth and declined with latitude The importance of accounting for this depth-latitude covariation in the distribution of marine fish demonstrated here, together with recent evidence obtained for deep-sea benthic macrofauna, points to depth and latitude as the main factors in the distribution of marine animals     

Patterns of diversity, depth range and body size among pelagic fishes along a gradient of depth

Studies of geographical patterns of diversity have focused largely on compiling and analysing data to evaluate alternative hypotheses for the near‐universal decrease in species richness from the equator to the poles. Valuable insights into the mechanisms that promote diversity can come from studies of other patterns, such as variation in species distributions with elevation in terrestrial systems or with depth in marine systems. To obtain such insights, we analysed and interpreted data on species diversity, depth of occurrence and body size of pelagic fishes along an oceanic depth gradient. We used a database on pelagic marine fishes native to the north‐east Pacific Ocean between 40°N and 50°N. We used data from the Pacific Rim Fisheries Program that were obtained from commercial, management and scientific surveys between 1999 and 2000. Depth of occurrence and maximum body length were used to assess the distributions of 409 species of pelagic fishes along a depth gradient from 0 to 8000 m. A presence–absence matrix was used to classify the depth range of each species into 100‐m intervals. Atmar & Patterson's (1995 ) software was used to quantify the degree of nestedness of species distributions. Pelagic fish species diversity decreased steeply with increasing depth; diversity peaked at less than 200 m and more than half of the species had mean depths of occurrence between 0 and 300 m. The distribution of species showed a very strong nested subset pattern along the depth gradient. Whereas species with narrow ranges were generally restricted to shallow waters, wide‐ranging species occurred from near the surface to great depths. The relationship between maximum body size and mean depth range differed between teleost and elasmobranch fishes: being positive for teleosts, but negative for elasmobranches. Results support hypotheses that some combination of high productivity and warm temperature promote high species diversity, and reject those that would attribute the pattern of species richness to the mid‐domain effect, habitat area, or environmental constancy. The data provided a clear example of Rapoport's rule, a negative correlation between average depth range and species diversity.     

长山列岛邻近海域鱼类群落种类组成和多样性时空变化

为了解黄、渤海生态系统交错带长山列岛邻近水域鱼类群落种类组成和多样性,根据2016年10月,2017年1月、5月及8月进行的鱼类资源底拖网调查数据,应用相对重要性指数、物种多样性指数、k优势度曲线等方法,研究了长山列岛邻近海域鱼类群落种类组成、物种多样性时空变化及其与环境因子的关系。结果表明: 该海域共计捕获鱼类77种,以温水性、底层、洄游性鱼类为主,优势种组成季节变化明显,春季、冬季主要优势种为黄鮟鱇,夏季以日本鲭、鳀等中上层鱼类为主。全年共计出现洄游性鱼类46种,季节间物种迁移指数均在100以上,其中秋季物种迁移指数最大。鱼类群落物种丰富度指数在春季最高、Shannon多样性指数和均匀度指数在秋季最高。夏季物种丰富度指数与表层水温呈显著负相关;冬季物种丰富度指数与水深、底层水温呈极显著正相关,Shannon多样性指数与底层水温呈极显著正相关。长山列岛邻近海域作为黄、渤海两大生态系统的交错带,鱼类群落表现出高物种多样性、洄游种多以及明显的时空异质性。     

Hierarchical analyses of genetic variation of samples from breeding and feeding grounds confirm the genetic partitioning of northwest Atlantic and South Atlantic populations of swordfish (Xiphias gladius L.)

In species with high migratory potential, the genetic signal revealing population differentiation is often obscured by population admixture. To our knowledge, the explicit comparison of genetic samples from known spawning and feeding areas has not been conducted for any highly migratory pelagic fish species. This study examines the geographic heterogeneity of swordfish mitochondrial DNA (mtDNA) lineages within the Atlantic Ocean using 330 base pairs of sequence of the control region from 480 individuals. Hierarchical analyses of sequence variation were conducted to test whether samples from areas identified as the corresponding spawning and feeding grounds for the northwest (NW) Atlantic (Caribbean and Georges Banks-US northeast) and the South Atlantic (Brazil-Uruguay and Gulf of Guinea), were more closely related to each other than to samples from any other region, including the Mediterranean Sea, the Indian Ocean, and the Pacific Ocean. Phylogeographic analyses reveal that swordfish mtDNA phylogeny is characterized by incomplete lineage sorting and secondary contact of two highly divergent clades. However, despite this complex phylogenetic signature, results from an analysis of nucleotide diversity and from an analysis of molecular variance (AMOVA) were for the most part concordant and indicate that NW Atlantic and South Atlantic swordfish belong to separate populations. The mtDNA distinctiveness of NW Atlantic and South Atlantic swordfish populations is indicative of philopatric behavior in swordfish towards breeding and feeding areas.     

Species range size distributions for some marine taxa in the Atlantic Ocean. Effect of latitude and depth

The range size distributions of 6643 species in ten different fish and invertebrate taxa dwelling in pelagic (latitudinal range sizes) and benthic (latitudinal and depth range sizes) habitats on both sides of the Atlantic Ocean (80°N−70°S) were studied. The objectives were to analyse: (1) the range size distribution patterns for the various taxa and whether they have right/left skewed or lognormal distributions; (2) the geographical species distributions, to ascertain whether the distribution ranges change with latitude (Rapoport's rule); and (3) the relationship between the depth ranges of benthic species and their maximum depth of occurrence and how depth range size distributions change with latitude. The pelagic taxa exhibited larger range sizes than did the benthic taxa, continental slope/rise species excepted. On the other hand, the boundaries between geographical provinces for both benthic taxa and pelagic taxa tended to occur in association with major oceanographic processes. The shape of the latitudinal range frequency distributions (LRFDs) of the pelagic organisms were distinctly left‐skewed, and the LRFDs for most taxa were significantly different from lognormal. There was no common pattern for the distributions of the benthic organisms, which were lognormal in Cephalopoda, Stomatopoda, and Crustacea Decapoda and tended to be left‐skewed and significantly different from lognormal in Pisces. The applicability of Rapoport's rule was not clearly inferable from the results, and the rule appears to be conditioned by the location of biogeographical boundaries and the endemism rate in the different biogeographical provinces. A clear increase in depth range size with maximum depth range was observable for benthic species, confirming previous studies. Species’ depth range distributions displayed a discernible latitudinal pattern, right‐skewed at high latitudes and left‐skewed at low latitudes. The location of biogeographical boundaries, and endemism rate by biogeographical province were considered to be the factors most useful in explaining species’ distribution patterns and their conformity or nonconformity to Rapoport's rule. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 80 , 437–455.     

Life histories of closely related amphidromous and non‐migratory fish species: a trade‐off between egg size and fecundity

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Rapid changes in fish community structure and habitat distribution following the precipitation of lake phosphorus with aluminium

1. Fish community structure and habitat distribution of the abundant species roach, perch and ruffe were studied in Lake Nordborg (Denmark) before (August 2006) and after (August 2007) aluminium treatment to reduce internal phosphorus loading. 2. Rapid changes in fish community structure, abundance and habitat distribution occurred following a decline in in‐lake phosphorus concentrations from 280 to 37 μg P L^?1 and an increase in Secchi depth transparency from 1.1 to 1.9 m (August). The proportion of perch in overnight gill net catches increased, whilst roach decreased, and the average weight of all key species increased. 3. The habitat distribution of perch and roach changed from a high proportion in the upper pelagic and littoral zones in 2006, towards enhanced proportions in the deeper pelagic and profundal zone in 2007. The abundance of large‐bodied zooplankton increased and the abundance of benthic invertebrates decreased in the same period, suggesting that the habitat shift was not induced by food limitation. 4. Ruffe shifted from the littoral and upper profundal zones towards the deep profundal zone, likely reflecting an increased predation risk in the littoral zone and better oxygen conditions in the deep profundal. 5. Our results indicate that enhanced risk of predation in the upper pelagic and the littoral zones and perhaps improved oxygen concentrations in the deeper profundal zone at decreasing turbidity are responsible for the observed habitat shift. The results indicate that fish respond rapidly to changes in nutrient state, both in terms of community structure and habitat use.     

Parasites as biological tags in population studies of demersal and pelagic fish species

Among the different techniques applied in a holistic approach for fish stock identification, the use of parasites as "biological tags" is becoming increasingly important. In this presentation, our recent studies on the use of some parasite species, identified by genetic markers, and the parasite/fauna composition, in stock identification of demersal (Merluccius merluccius), small pelagic (Trachurus trachurus), and large pelagic fish species (Xiphias gladius) are reviewed. Different species of Anisakis and Hysterothylacium were genetically identified by the application of genetic (allozyme) markers. Statistically significant differences in the spatial distribution of distinct species of Anisakis were found in the fish considered. As to the species of Hysterothylacium genetically detected, different relative proportions were detected in several Mediterranean and Atlantic samples of swordfish (X. gladius). This study demonstrates the potential value of these anisakid nematodes, at both larval and adult stages, as "biological tags" for these fish species in European waters.     

Global phylogeography of the dolphinfish (Coryphaena hippurus): the influence of large effective population size and recent dispersal on the divergence of a marine pelagic cosmopolitan species

Pelagic fish that are distributed circumtropically are characterised by a low population structure level as a result of a high capacity for dispersion and large population sizes. Nevertheless, historical and contemporary processes, including past demographic and/or range expansions, secondary contact, dispersal, gene flow, and the achievement of large effective population sizes, may play a part in the detection of divergence signals, especially in the case of tropical pelagic species, whose distribution range depends strongly on the sea surface temperature. The connectivity and historical demography of Atlantic, Indian, Pacific and Mediterranean populations of dolphinfish (Coryphaena hippurus) was studied using partial sequences of the mitochondrial DNA NADH dehydrogenase subunit 1 (ND1). AMOVA analyses revealed significant inter-oceanic divergence with three phylogroups located in the Indo-Pacific, Eastern Atlantic, and Mediterranean Sea, the last one being the most divergent. However, it was not possible to clearly observe any genetic differentiation between the Indo-Pacific and Atlantic populations, as has been reported for most tropical pelagic species of tuna and billfishes. This supports the assumption of recent dispersal among basins facilitated by the actual continuous distribution of dolphinfish populations. Moreover, the lack of a divergence signal for populations separated by the Panamanian Isthmus reveals that genetic drift does not exert a strong influence on tropical pelagic species with large effective population sizes.     

Warming shelf seas drive the subtropicalization of European pelagic fish communities

Pelagic fishes are among the most ecologically and economically important fish species in European seas. In principle, these pelagic fishes have potential to demonstrate rapid abundance and distribution shifts in response to climatic variability due to their high adult motility, planktonic larval stages, and low dependence on benthic habitat for food or shelter during their life histories. Here, we provide evidence of substantial climate‐driven changes to the structure of pelagic fish communities in European shelf seas. We investigated the patterns of species‐level change using catch records from 57 870 fisheries‐independent survey trawls from across European continental shelf region between 1965 and 2012. We analysed changes in the distribution and rate of occurrence of the six most common species, and observed a strong subtropicalization of the North Sea and Baltic Sea assemblages. These areas have shifted away from cold‐water assemblages typically characterized by Atlantic herring and European sprat from the 1960s to 1980s, to warmer‐water assemblages including Atlantic mackerel, Atlantic horse mackerel, European pilchard and European anchovy from the 1990s onwards. We next investigated if warming sea temperatures have forced these changes using temporally comprehensive data from the North Sea region. Our models indicated the primary driver of change in these species has been sea surface temperatures in all cases. Together, these analyses highlight how individual species responses have combined to result in a dramatic subtropicalization of the pelagic fish assemblage of the European continental shelf.     

Spatial Scale,Means and Gradients of Hydrographic Variables Define Pelagic Seascapes of Bluefin and Bullet Tuna Spawning Distribution

Seascape ecology is an emerging discipline focused on understanding how features of the marine habitat influence the spatial distribution of marine species. However, there is still a gap in the development of concepts and techniques for its application in the marine pelagic realm, where there are no clear boundaries delimitating habitats. Here we demonstrate that pelagic seascape metrics defined as a combination of hydrographic variables and their spatial gradients calculated at an appropriate spatial scale, improve our ability to model pelagic fish distribution. We apply the analysis to study the spawning locations of two tuna species: Atlantic bluefin and bullet tuna. These two species represent a gradient in life history strategies. Bluefin tuna has a large body size and is a long-distant migrant, while bullet tuna has a small body size and lives year-round in coastal waters within the Mediterranean Sea. The results show that the models performance incorporating the proposed seascape metrics increases significantly when compared with models that do not consider these metrics. This improvement is more important for Atlantic bluefin, whose spawning ecology is dependent on the local oceanographic scenario, than it is for bullet tuna, which is less influenced by the hydrographic conditions. Our study advances our understanding of how species perceive their habitat and confirms that the spatial scale at which the seascape metrics provide information is related to the spawning ecology and life history strategy of each species.     

Environmental correlates of population differentiation in Atlantic herring

The marine environment is characterized by few physical barriers, and pelagic fishes commonly show high migratory potential and low, albeit in some cases statistically significant, levels of genetic divergence in neutral genetic marker analyses. However, it is not clear whether low levels of differentiation reflect spatially separated populations experiencing gene flow or shallow population histories coupled with limited random genetic drift in large, demographically isolated populations undergoing independent evolutionary processes. Using information for nine microsatellite loci in a total of 1951 fish, we analyzed genetic differentiation among Atlantic herring from eleven spawning locations distributed along a longitudinal gradient from the North Sea to the Western Baltic. Overall genetic differentiation was low (theta = 0.008) but statistically significant. The area is characterized by a dramatic shift in hydrography from the highly saline and temperature stable North Sea to the brackish Baltic Sea, where temperatures show high annual variation. We used two different methods, a novel computational geometric approach and partial Mantel correlation analysis coupled with detailed environmental information from spawning locations to show that patterns of reproductive isolation covaried with salinity differences among spawning locations, independent of their geographical distance. We show that reproductive isolation can be maintained in marine fish populations exhibiting substantial mixing during larval and adult life stages. Analyses incorporating genetic, spatial, and environmental parameters indicated that isolating mechanisms are associated with the specific salinity conditions on spawning locations.     

Year-to-year variation in larval fish assemblages of the Southern North Sea

In order to test the temporal stability within and the reproducibility of larval fish assemblages between years, the larval fish assemblage at Helgoland Roads, North Sea (NE Atlantic) was quantitatively sampled almost daily from January 2003 to December 2005. The survey resulted in a total of 462 samples containing 50,632 larval fish of at least 42 taxa. In winter the larval fish assemblage was mainly dominated by larvae emerging from demersal eggs. This changed gradually to larvae hatching from pelagic eggs. Larvae from pelagic eggs dominated the ichthyoplankton assemblage in summer. A remarkably stable seasonality in terms of dominance patterns with recurring, season-specific fish assemblages was observed over the 3 years, despite substantial variation in environmental conditions such as a temperature difference of almost 20°C between summer and winter. The lesser sandeel (Ammodytes marinus), was the only species which showed significant fluctuations in abundance between the years. After removal of this species from the analysis, the dominance patterns of the remaining fish species were almost identical between years.