Manuel Hidalgo

Abstract: Fisheries ecologists traditionally aimed at disentangling climate and fishing effects from the population dynamics of exploited marine fish stocks. However, recent studies have shown that internal characteristics and external forcing (climate and exploitation) have interactive rather than additive effects. Thought most of these studies explored how demographic truncation induced by exploitation affected the response of recruitment to climate, identifying a general pattern revealed to be difficult as interactions are often case-specific. Here we compared five exploited stocks of European hake Merluccius merluccius from the Atlantic Ocean and Mediterranean Sea to investigate how the interaction between internal characteristics and external forces affect the variability of the population growth rate and their consequences on recruitment. Our results show that demographic truncation induces a novel population scenario in which the growth rate is maximized when the reproductive stock is younger and less diverse. This scenario is shaped by the climate variability and the fishing pattern. The population growth rate becomes more dependent on the maturation schedule and less on the survival rates. The consequences for the recruitment dynamics are twofold; the effect of density-dependent regulatory processes decreases while the effect of the density-independent drivers increases. Our study shows that the interaction between internal characteristics and external forces changes across geographic locations according to 1) the importance of demographic truncation, 2) the influence of the climate on the regional hydrography and 3) the spatiotemporal heterogeneity of the physical environment to which fish life history is adapted.

Abstract: Biological processes and physical oceanography are often integrated in numerical modelling of marine fish larvae, but rarely in statistical analyses of spatio-temporal observation data. Here, we examine the relative contribution of inter-annual variability in spawner distribution, advection by ocean currents, hydrography and climate in modifying observed distribution patterns of cod larvae in the Lofoten–Barents Sea. By integrating predictions from a particle-tracking model into a spatially explicit statistical analysis, the effects of advection and the timing and locations of spawning are accounted for. The analysis also includes other environmental factors: temperature, salinity, a convergence index and a climate threshold determined by the North Atlantic Oscillation (NAO). We found that the spatial pattern of larvae changed over the two climate periods, being more upstream in low NAO years. We also demonstrate that spawning distribution and ocean circulation are the main factors shaping this distribution, while temperature effects are different between climate periods, probably due to a different spatial overlap of the fish larvae and their prey, and the consequent effect on the spatial pattern of larval survival. Our new methodological approach combines numerical and statistical modelling to draw robust inferences from observed distributions and will be of general interest for studies of many marine fish species.

Abstract: The synergistic effects of fishing, climate and internal dynamics on population fluctuations are poorly understood due to the complexity of these interactions. In this paper, we combine time series analysis and simulations to investigate the long-term dynamics of an overexploited population in the Mediterranean Sea, and its link with both fishing-induced demographic changes and hydroclimatic variability. We show that the cyclicity of the catch per unit of effort (CPUE) of European hake Merluccius merluccius (EH) vanished in the 1980s, while the correlation between the CPUE and a local environmental index increased. Using simulations, we then show that the cyclicity observed in the EH biomass before the 1980s can have an internal origin, while that its disappearance could be due to the fishing-induced erosion of the age structure. Our results suggest that fishing can trigger a switch from internally generated to externally forced population fluctuations, the latter being characterised by an increasing dependency of the population on recruitment and ultimately on environmental variability. Hydroclimatic modifications occurring in the Mediterranean in the early 1980s could have enhanced these changes by leading to a mismatch between early life stages of EH and favorable environmental conditions. Our conclusions underline the key effect of the interaction between exploitation and climate on the dynamics of EH and its important consequences for management and conservation.

Abstract: Accumulating evidence shows that environmental fluctuations and exploitation jointly affect marine fish populations, and understanding their interaction is a key issue for fisheries ecology. In particular, it has been proposed that age truncation induced by fisheries exploitation may increase the population's sensitivity to climate. In this study, we use unique long-term abundance data for the Northeast Arctic stock of cod (Gadus morhua) and the Norwegian Spring-Spawning stock of herring (Clupea harengus), which we analyze using techniques based on age-structured population matrices. After identifying time periods with different age distributions in the spawning stock, we use linear models to quantify the relative effect of exploitation and temperature on the population growth rates. For the two populations, age truncation was found to be associated with an increasing importance of temperature and a relatively decreasing importance of exploitation, while the population growth rate became increasingly sensitive to recruitment variations. The results suggested that the removal of older age classes reduced the buffering capacity of the population, thereby making the population growth rate more dependent on recruitment than adult survival and increasing the effect of environmental fluctuations. Age structure appeared as a key characteristic that can affect the response of fish stocks to climate variations and its consequences may be of key importance for conservation and management.

Abstract: Basic information on the distribution and habitat preferences of ecologically important species is essential for their management and protection. This study focuses on the depth related trends and the geographic patterns that shape the community of the elasmobranch species in the Balearic Islands (Mediterranean Sea) using data collected from 2001 to 2009. Non-metric Multi-Dimensional Scaling (MDS) ordination was used to detect zonation patterns in the community. Generalized Additive Models (GAMs) were applied to analyse spatial and temporal variation in elasmobranch community descriptors (abundance, biomass, mean fish weight, number of species and diversity), as well as the abundance and mean length of the four individual species (S. canicula, G. melastomus, R. clavata, R. miraletus). Depth was the main factor determining the assemblage composition, and the MDS analysis identified four main groups with 60% of the similarity found to correspond to the continental shelf, shelf break, upper slope and middle slope of the surveyed area. GAM analysis identified spatial patterns that were independent of the bathymetric distribution preference. Although depth was a strong predictor for all the analyses performed, the geographic variation in the elasmobranch abundance was also important. The results also show a reduction in the mean length of the elasmobranch species in the areas with high fishing intensity. Our study evidences a clear spatial segregation of the main species throughout the ontogeny because the geographic and bathymetric effects were highly size dependent, with clear differences between the bathymetric distributions of juveniles and adults but no clear spatial overlapping. This study sheds new light on the spatial distribution of the elasmobranch species off the Balearic Islands, which is essential information for protecting marine organisms along with their habitats and promoting ecosystem based

Abstract: Seasonal pattern of fish recruitment can vary over years, and the coupling of such pattern with favourable trophic-environmental conditions may trigger non-stationary and non-linear effects on the yearly recruitment variability. The relative contribution of intra-annual discrete recruitment events (REs) to recruitment is often the basis of the inter-annual fluctuations in abundance and biomass of exploited fish populations which are based on a small number of age classes. In this study, REs of European hake (Merluccius merluccius L.) were identified in the north and south of the Balearic Islands (NW Mediterranean) during the 2004 recruitment season and investigated for growth and physiological and morphological condition. Five REs were identified with different contributions in abundance to the annual recruitment in each area coming from different intra-annual spawning batches. Growth rates were similar between the REs with the exception of the last event of the season. Conversely, large differences in morphological and physiological condition were observed between REs and were due to: different environmental-trophic scenarios encountered, and ontogenetic changes related to the recruitment to the shelf-break grounds (ca. 11 cm, 5–6 months of life) which triggered a trade-off in the energy allocation strategy. The spatial-temporal variability in the physiological condition among REs at the beginning of the recruitment season might be explained in the basis on a ‘condition expanded match–mismatch hypothesis’. We advance that individuals recruiting in the north, which were in better condition and temporally and spatially differenced from the south, have most probably hatched in separated locations with different food availability for larvae and early life stages. We propose that hake recruiting to the north and south of the island are from different origins, mainland and the islands respectively, and have benefited from different environmental conditions. This results in a different condition but not in different growth rates.

Abstract: In 1980 and 1995, the European hake (Merluccius merluccius) population off the Balearic Islands (northwestern Mediterranean) underwent changes at both the individual level and the population level. There was a sharp decrease in abundance that coincided with a change in the seasonal catch-per-unit-effort pattern in 1980. A new population scenario emerged after 1980 characterized by an increase in the intrinsic growth rate and a decrease in carrying capacity; however, catchability remained the same. An age-structure truncation could have caused these changes, making the population more dependent on year-to-year recruitment. A change in size structure also occurred in 1995, which was evidenced by a sudden decrease in the mean length-at-age (L) and abundance of recruits and a change in the density-dependent effect on recruits. As the Mediterranean trawl fishery mainly harvests recruits and juveniles and fishery harvesting induces adaptive changes in life history traits, the sharp decrease in L of recruits could be explained as a growth reduction due to the selective pressure to stay under mesh size for longer and thus maximize survival until reproduction. These individual and population transitions explain the changes in the response to the environmental forcing observed in the European hake off the Balearic Islands.

Abstract: Short spatio–temporal variations in the feeding intensity and the diet of the European hake, Merluccius merluccius, together with the abundance of their potential prey were studied between August 2003 and June 2004 at two locations, northwest (Sóller) and south (Cabrera), off the island of Mallorca (Balearic Islands, Western Mediterranean) at depths between 150 and 750 m. The two areas present different oceanographic conditions. Hake was mainly distributed along the shelf-slope break and the upper slope (between 166 and 350 m) where recruits (TL<18 cm) were dominant. The hake's diet varied as a function of size. Recruits fed mainly on micronektonic prey, and the diet was influenced primarily by seasonality, with two dietary patterns (identified by MDS analyses) corresponding to August–September 2003 (summer) and to November 2003/February–April 2004 (autumn–winter). The summer pattern was consistent with a thermally stratified water column, while November and April were consistent with homogenized temperature and salinity throughout all the water column. The main prey of recruits were the euphausiid Meganyctiphanes norvegica and the midwater fish Maurolicus muelleri in autumn–winter and myctophids (mainly Ceratoscopelus maderensis) in summer. In contrast to recruits, the geographic factor (NW vs. S) was the main factor influencing the diets of post-recruits (TL between 18 and 21.9 cm) and adults (TL ≥ 22 cm). Hake recruits (and to a lesser extent post-recruits) and their preferred prey occupied different depth ranges during daylight periods. Meganyctiphanes norvegica and Ceratoscopelus maderensis were, for instance, distributed as much as 500 m deeper than hake that had eaten them. All these trends were especially obvious at NW, an area with a more abrupt slope and with a greater influence by northern winter intermediate water (WIW) inflow in early spring than the S area. These factors probably enhanced micronekton aggregation in April, when feeding intensity (stomach fullness) increased among recruits and post-recruits only at NW. All these factors may have a crucial role in the diet, distribution and probably recruitment success of small hake. Biological factors were also important in trophic shifts in the diet and feeding of hake. Multi-linear regression models pointed to a trend of higher fullness with higher hepato-somatic index (HSI). Therefore greater food consumption by hake may enhance its metabolic condition. Within the framework of shelf-break and slope ecology, we show how the ‘boundary’ mesopelagic community inhabiting the middle slope sustains the trophic requirements of hake, a species distributed at shallower depths along the shelf-slope break. Mesopelagic euphausiids and myctophids are often found in the diets of shelf-break fish. Because the boundary mesopelagic community is distributed worldwide, the high levels of fish biomass often found at shelf-slope breaks could be sustained trophically by deeper, offshore mesopelagic communities, an inverse energy transfer from deep to shallow-water marine ecosystems.

Abstract: The aim of the present paper is to study the relationships between some climatic indices and parental stock, recruitment and accessibility to trawl fishery of hake (Merluccius merluccius) and red shrimp (Aristeus antennatus) off Balearic Islands (western Mediterranean). Available annual catch per unit effort, recruitment and spawning stock biomass have been used as biological data. As environmental data, the mesoscale IDEA index and the large-scale North Atlantic Oscillation (NAO) and Mediterranean Oscillation (MO) indices have been used. To analyze possible links between these indices with the population dynamics of demersal resources, two non-linear approaches have been applied: (i) stock–recruitment relationships from Ricker and Beverton–Holt models, by sequentially incorporating environment factors; (ii) generalized additive modelling, both classical general and threshold non-additive models were considered. The latter simulate an abrupt change in explicative variables across different phases (time periods or climatic index values). The results have shown that two oceanographic scenarios around the Balearic Islands, associated with macro and meso-scale climate regimes, can influence the population dynamics of hake and red shrimp. This is especially true for recruitment, which seems to be enhanced during lowNAOand IDEAindices periods.During these periods, colder-than-normal winters generate high amounts of coldWestern Mediterranean IntermediateWaters (WIW) in the Gulf of Lions, which flow southwards and reach the Balearic Islands channels in spring, increasing the productivity in the area. This oceanographic scenario could also be favourable to the distribution of hake on the fishing grounds where the trawl fleet targets this species, increasing its accessibility to the fishery. Both spawning stock and abundance of red shrimp seems to be also enhanced by high MO index periods, which could reflect the increased presence of the saline and warm Levantine IntermediateWaters (LIW) in the study area, extending over the fishing grounds of this species. The proposed interactions can be useful to assess and manage these important demersal resources.

Abstract: The spatial and temporal variations of deep-sea megafaunal assemblages from the western Mediterranean are analysed in the present paper. The assemblages from two locations of the Balearic Islands situated 120 km apart were compared using data collected seasonally on a bathymetric stratum covering the 150–750 m depth range during six bottom-trawl surveys. The assemblage structure, in terms of species composition, species dominance and population sizes, was differentially affected by the spatio-temporal variables analysed (depth, location and fishing period). Although depth was the main factor determining the assemblage composition, the differences obtained between the two locations were also relevant. On the upper slope these between-location differences in the dynamics of megafaunal assemblages were found to be related to the effect of fishing exploitation. Population size-based metrics and biomass spectra were good predictors of meso-scale fishing effects, and were mainly reflected by elasmobranchs and demersal teleosts. Nevertheless, the effects of fishing depended on the species considered. Two dominant large-sized fish species found on the upper slope in both localities, Galeus melastomus and Phycis blennoides, had higher biomass values associated with lower fishing effort. Although the mean body weight (MBW) of both species and also the mean maximum body weight (MMBW) of G. melastomus agreed with this pattern, the P. blennoides did not. This last case could be indicative of natural size-trends such as the bigger– deeper phenomenon which refers to the displacement of large individuals towards the deeper limit of their bathymetric distribution, beyond the maximum depth sampled in this study for this species. By contrast, the target species of the upper slope fishery, the red shrimp Aristeus antennatus, was not negatively affected by the direct impact of fishing activity and other environmental factors, such as the presence of specific water masses could also be important.

Abstract: Oxygen (δ18O) and carbon (δ13C) isotope ratios in sagittal otoliths were analysed in recruits and juveniles of European hake Merluccius merluccius L. caught in the northwest (Sóller, SO) and south (Cabrera, CA) of the Island of Mallorca (western Mediterranean) over 2 consecutive years (2003 and 2004). The analytical method used allowed data to be gathered on both environmental and trophic conditions experienced by fish during the pelagic early life stages (registered in the composition of the inner part of the otolith, core area) and the demersal aggregations of recruits (at the otolith edge). Results on the seasonal variation of oxygen isotope signatures at the edge of fish otoliths captured in SO and CA indicated different vertical migration behaviours of hake between the bottom and the thermocline. δ13C showed a clear ontogenetic pattern between the core area and the edge, confirming the differences in diet from the early stages to post-recruits. The seasonal analysis of δ13C data from the otolith edge indicated that fish from the early stages in June 2003 encountered poorer trophic resources, which resulted in poor fish condition and subsequent poorer recruitment. The estimated temperature from the core area of otoliths showed lower temperature regimes in the 2002 hatching season compared to the 2003 hatching season, which could be a possible explanation for the observed differences in success of subsequent recruitment.

Abstract: This study evaluates the link between the recruitment process of European hake (Merluccius merluccius L.) of the Balearic Islands (western Mediterranean) and the environmental and physiological conditions. Spatio-temporal variation of abundance and condition of fish were evaluated at two locations each with different oceanographic conditions, one in the north (Sóller, SO) and another in the south (Cabrera, CA) of Mallorca Island. Environmental variables explored were hydrography, sediment characteristics, phytoplankton pigment concentration (ppc) and the trophic resources of hake. Individuals were divided in three life stages: recruits, post-recruits and young adults. Hepatosomatic index (HSI), relative condition index (Kn), gonadosomatic index (GSI) and fullness index (FI) were analysed for the three life stages. Recruitment starts in February with the incorporation of smaller hakes, and it can be followed through spring and early summer with a peak in April. However, some spatial heterogeneity in the recruitment process has been found between north and south of the Island. The main pulse of recruitment occurred at a different time in the two areas. Spatial heterogeneity was also consistent with the condition of hake recruits, with higher values of Kn and HSI at SO than at CA. Maximum values of Kn were found in February at SO and in April at CA, coinciding with the start of the different recruitment pulses to the fishing grounds. Post-recruits and young adults also showed higher condition at SO than at CA. The arrival in spring of the Western Winter Intermediate Waters (WIW) drives the spatial-temporal variation in abundance and condition of hake. Ppc was highly correlated with recruit abundance with a time lag of two months, while for post-recruits the time lag was three months. The observed differences in the condition of hake between areas could be a consequence of the fact that the waters to the north of Mallorca are comparatively more under the seasonal influence of WIW which is formed in more productive areas. Thus, this study characterises the short temporal and spatial variability in the hake recruitment process off the Balearic Islands, both in terms of abundance and fish condition. This pattern is explained on the basis of the mesoscale environmental variability observed between north and south of Mallorca and the ecological adaptive strategy of recruiting in the optimal environmental season.