Trophic indicators in fisheries: a call for re-evaluation.

Mean trophic level (MTL) of landings and primary production required (PPR) by fisheries are increasingly used in the assessment of sustainability in fisheries. However, in their present form, MTL and PPR are prone to misinterpretation. We show that it is important to account for actual catch data, define an appropriate historical and spatial domain, and carefully consider the effects of fisheries management, based on results from a case study of Swedish fisheries during the past century.

Non-additive and non-stationary properties in the spatial distribution of a large marine fish population.

Density-independent and density-dependent variables both affect the spatial distributions of species. However, their effects are often separately addressed using different analytical techniques. We apply a spatially explicit regression framework that incorporates localized, interactive and threshold effects of both density-independent (water temperature) and density-dependent (population abundance) variables, to study the spatial distribution of a well-monitored flatfish population in the eastern Bering Sea. Results indicate that when population biomass was beyond a threshold a further increase in biomass-promoted habitat expansion in a non-additive fashion with water temperature. In contrast, during years of low population size, habitat occupancy was affected positively only by water temperature. These results reveal the spatial signature of intraspecific abundance distribution relationships as well as the non-additive and non-stationary responses of species spatial dynamics. Furthermore, these results underscore the importance of implementing analytical techniques that can simultaneously account for density-dependent and density-independent sources of variability when studying geographical distribution patterns.

Trophic cascades promote threshold-like shifts in pelagic marine ecosystems.

Fisheries can have a large impact on marine ecosystems, because the effects of removing large predatory fish may cascade down the food web. The implications of these cascading processes on system functioning and resilience remain a source of intense scientific debate. By using field data covering a 30-year period, we show for the Baltic Sea that the underlying mechanisms of trophic cascades produced a shift in ecosystem functioning after the collapse of the top predator cod. We identified an ecological threshold, corresponding to a planktivore abundance of approximately 17 x 10(10) individuals, that separates 2 ecosystem configurations in which zooplankton dynamics are driven by either hydroclimatic forces or predation pressure. Abundances of the planktivore sprat above the threshold decouple zooplankton dynamics from hydrological circumstances. The current strong regulation by sprat of the feeding resources for larval cod may hinder cod recovery and the return of the ecosystem to a prior state. This calls for the inclusion of a food web perspective in management decisions.

Ontogenetic and sex-specific differences in density-dependent habitat selection of a marine fish population.

The spatial dynamics of species are the result of complex interactions between density-independent and density-dependent sources of variability. Disentangling these two sources of variability has challenged ecologists working in both terrestrial and aquatic ecosystems. Using a novel spatially explicit statistical model, we tested for the presence of density-independent and density-dependent habitat selection in yellowfin sole (Limanda aspera) in the eastern Bering Sea. We found specificities in the density-dependent processes operating across ontogeny and particularly with gender. Density-dependent habitat expansion occurred primarily in females, and to a lesser degree in males. These patterns were especially evident in adult stages, while juvenile stages of both sexes exhibited a mix of different dynamics. Association of yellowfin sole with substrate type also varied by sex and to a lesser degree with size, with large females distributed over a wider range of substrates than males. Moreover, yellowfin sole expanded northward as cold subsurface waters retracted in summer, suggesting high sensitivity to arctic warming. Our findings illustrate how marginal habitats can play an important role in buffering density-dependent habitat expansion, with direct implications for resource management. Our spatially explicit modeling approach is effective in evaluating density-dependent spatial dynamics, and can easily be used to test similar hypotheses from a variety of aquatic and terrestrial ecosystems.

Model uncertainty and simulated multispecies fisheries management advice in the Baltic Sea.

Different ecosystem models often provide contrasting predictions (model uncertainty), which is perceived to be a major challenge impeding their use to support ecosystem-based fisheries management (EBFM). The focus of this manuscript is to examine the extent of model disagreements which could impact management advice for EBFM in the central Baltic Sea. We compare how much three models (EwE, Gadget and a multispecies stock production model) differ in 1) their estimates of fishing mortality rates (Fs) satisfying alternative hypothetical management scenario objectives and 2) the outcomes of those scenarios in terms of performance indicators (spawning stock biomasses, catches, profits). Uncertainty in future environmental conditions affecting fish was taken into account by considering two seal population growth scenarios and two nutrient load scenarios. Differences in the development of the stocks, yields and profits exist among the models but the general patterns are also sufficiently similar to appear promising in the context of strategic fishery advice. Thus, we suggest that disagreements among the ecosystem models will not impede their use for providing strategic advice on how to reach management objectives that go beyond the traditional maximum yield targets and for informing on the potential consequences of pursuing such objectives. This is especially true for scenarios aiming at exploiting forage fish sprat and herring, for which the agreement was the largest among our models. However, the quantitative response to altering fishing pressure differed among models. This was due to the diverse environmental covariates and the different number of trophic relationships and their functional forms considered in the models. This suggests that ecosystem models can be used to provide quantitative advice only after more targeted research is conducted to gain a deeper understanding into the relationship between trophic links and fish population dynamics in the Baltic Sea.

Classifying grey seal behaviour in relation to environmental variability and commercial fishing activity - a multivariate hidden Markov model.

Classifying movement behaviour of marine predators in relation to anthropogenic activity and environmental conditions is important to guide marine conservation. We studied the relationship between grey seal (Halichoerus grypus) behaviour and environmental variability in the southwestern Baltic Sea where seal-fishery conflicts are increasing. We used multiple environmental covariates and proximity to active fishing nets within a multivariate hidden Markov model (HMM) to quantify changes in movement behaviour of grey seals while at sea. Dive depth, dive duration, surface duration, horizontal displacement, and turning angle were used to identify travelling, resting and foraging states. The likelihood of seals foraging increased in deeper, colder, more saline waters, which are sites with increased primary productivity and possibly prey densities. Proximity to active fishing net also had a pronounced effect on state occupancy. The probability of seals foraging was highest <5 km from active fishing nets (51%) and decreased as distance to nets increased. However, seals used sites <5 km from active fishing nets only 3% of their time at sea highlighting an important temporal dimension in seal-fishery interactions. By coupling high-resolution oceanographic, fisheries, and grey seal movement data, our study provides a scientific basis for designing management strategies that satisfy ecological and socioeconomic demands on marine ecosystems.

Spatio-temporal dynamics of a fish predator: Density-dependent and hydrographic effects on Baltic Sea cod population.

Understanding the mechanisms of spatial population dynamics is crucial for the successful management of exploited species and ecosystems. However, the underlying mechanisms of spatial distribution are generally complex due to the concurrent forcing of both density-dependent species interactions and density-independent environmental factors. Despite the high economic value and central ecological importance of cod in the Baltic Sea, the drivers of its spatio-temporal population dynamics have not been analytically investigated so far. In this paper, we used an extensive trawl survey dataset in combination with environmental data to investigate the spatial dynamics of the distribution of the Eastern Baltic cod during the past three decades using Generalized Additive Models. The results showed that adult cod distribution was mainly affected by cod population size, and to a minor degree by small-scale hydrological factors and the extent of suitable reproductive areas. As population size decreases, the cod population concentrates to the southern part of the Baltic Sea, where the preferred more marine environment conditions are encountered. Using the fitted models, we predicted the Baltic cod distribution back to the 1970s and a temporal index of cod spatial occupation was developed. Our study will contribute to the management and conservation of this important resource and of the ecosystem where it occurs, by showing the forces shaping its spatial distribution and therefore the potential response of the population to future exploitation and environmental changes.

Large-Scale Spatio-Temporal Patterns of Mediterranean Cephalopod Diversity.

Species diversity is widely recognized as an important trait of ecosystems' functioning and resilience. Understanding the causes of diversity patterns and their interaction with the environmental conditions is essential in order to effectively assess and preserve existing diversity. While diversity patterns of most recurrent groups such as fish are commonly studied, other important taxa such as cephalopods have received less attention. In this work we present spatio-temporal trends of cephalopod diversity across the entire Mediterranean Sea during the last 19 years, analysing data from the annual bottom trawl survey MEDITS conducted by 5 different Mediterranean countries using standardized gears and sampling protocols. The influence of local and regional environmental variability in different Mediterranean regions is analysed applying generalized additive models, using species richness and the Shannon Wiener index as diversity descriptors. While the western basin showed a high diversity, our analyses do not support a steady eastward decrease of diversity as proposed in some previous studies. Instead, high Shannon diversity was also found in the Adriatic and Aegean Seas, and high species richness in the eastern Ionian Sea. Overall diversity did not show any consistent trend over the last two decades. Except in the Adriatic Sea, diversity showed a hump-shaped trend with depth in all regions, being highest between 200-400 m depth. Our results indicate that high Chlorophyll a concentrations and warmer temperatures seem to enhance species diversity, and the influence of these parameters is stronger for richness than for Shannon diversity.

Density-dependence in space and time: opposite synchronous variations in population distribution and body condition in the Baltic Sea sprat (Sprattus sprattus) over three decades.

Spatio-temporal density-dependent processes are crucial regulatory factors for natural populations. However, there is a lack of studies addressing spatial density-dependence in fish growth. A previous investigation has suggested spatio-temporal density-dependence in body condition of Baltic sprat. Here, we used different techniques, such as centre of gravity, distance, and homogeneity indices, to better characterize the spatial and temporal variations in sprat density and body condition in the Baltic Proper. Our results evidenced a negative spatio-temporal co-variation between the centres of gravity of density and maximum condition. In the 1980s-early 1990s both centres were located in the middle of the Baltic Proper. From the mid 1990s the centres progressively separated in space, as the sprat population moved towards the north-eastern Baltic Proper, and the centre of maximum condition towards the south-western areas. Moreover, at low abundances, sprat density and condition were homogeneously distributed in space, whereas at high abundances both density and condition showed pronounced geographical gradients. The ecological processes potentially explaining the observed patterns were discussed in the light of the Ideal Free Distribution theory. We provide evidence that the shift in the spatial distribution of cod, the main predator of sprat, has been the main factor triggering the overall spatial changes in sprat density, and thus condition, during the past thirty years. The spatial indices shown here, synthesizing the spatio-temporal patterns of fish distribution, can support the implementation of the EU Marine Strategy Framework Directive.

Multi-annual fluctuations in reconstructed historical time-series of a European lobster (Homarus gammarus) population disappear at increased exploitation levels.

Through the history of ecology, fluctuations of populations have been a dominating topic, and endogenous causes of fluctuations and oscillations have been recognized and studied for more than 80 years. Here we analyzed an historical dataset, covering more than 130 years, of European lobster (Homarus gammarus) catches. The data shows periodic fluctuations, which are first dampened and then disappear over time. The disappearance of the periodicity coincided with a substantial increase in fishing effort and the oscillations have not reappeared in the time series. The shifting baseline syndrome has changed our perception of not only the status of the stock, but also the regulating pressures. We describe the transition of a naturally regulated lobster population into a heavily exploited fisheries controlled stock. This is shown by the incorporation of environmental and endogenous processes in generalized additive models, autocorrelation functions and periodicity analyses of time-series.

Fishery discards: factors affecting their variability within a demersal trawl fishery.

Discards represent one of the most important issues within current commercial fishing. It occurs for a range of reasons and is influenced by an even more complex array of factors. We address this issue by examining the data collected within the Danish discard observer program and describe the factors that influence discarding within the Danish Kattegat demersal fleet over the period 1997 to 2008. Generalised additive models were used to assess how discards of the 3 main target species, Norway lobster, cod and plaice, and their subcomponents (under and over minimum landings size) are influenced by important factors and their potential relevance to management. Our results show that discards are influenced by a range of different factors that are different for each species and portion of discards. We argue that knowledge about the factors influential to discarding and their use in relation to potential mitigation measures are essential for future fisheries management strategies.

Morphological variation in the seahorse vertebral system / Variaciones morfológicas en el sistema vertebral del caballo de mar

The vertebral system in Hippocampus hippocampus is highly specialised because of the vertical locomotion and tail prehensility. The vertebral elements represent a special case of morphological changes, being the metameric structures organised along a natural functional series. We investigated the shape changes along the vertebral spine in H. hippocampus through geometric morphometrics, in order to describe functional and structural patterns. Actually, the dorso-ventral tail bending ability in the genus Hippocampus is one of the most impressive morphological modifications in the evolutionary history of fishes. Vertebrae were analysed using a 2D configuration from the left lateral view. The variation along the vertebral series suggests the identification of cervical, abdominal, dorsal, and caudal groups. The first three (cervical) elements and the lOth (supra-dorsal) structure show peculiar morphologies and local adaptations, associated with neck angulation and dorsal fin muscles, respectively. The vertebral size decreases from the anteriormost element backward, with some local variation at the dorsal area. Major changes are related to allometric variation at the neural región. The caudal elements are characterised by a marked size decrease, with consequent allometric shape changes involving the rotation of the posterior vertebral opening. This allometric trajectory leads to a natural ventral bending of the tail, promoting its prehensile function. This morphological survey suggests an interesting structural network between posture, locomotion, and vertebral anatomy, underlying the main functional changes in the Hippocampus biomechanics. Geometric morphometrics is rather suitable to approach metameric studies in terms of serial variation and functional adaptations.

El sistema vertebral en Hippocampus hippocampus es altamente especializado, debido a la locomoción vertical y la cola prehensible. Los elementos vertebrales representan un caso especial de cambios morfológicos, siendo estructuras metaméricas organizadas a lo largo de una serie funcional natural. Se investigaron los cambios en la forma, a lo largo de la columna vertebral en H. hippocampus a través de morfometría geométrica, con el fin de describir sus patrones funcionales y estructurales. Actualmente, la capacidad de flexión dorso-ventral de la cola en el género Hippocampus es una de las más impresionantes modificaciones morfológicas en la historia evolutiva de los peces. Las vértebras se analizaron mediante una configuración 2D de la vista lateral izquierda. La variación a lo largo de la serie vertebral sugiere la identificación de grupos cervicales, abdominales, dorsales, caudales. Los tres primeros elementos (cervicales) y la 10ª (supra-dorsal) muestran una estructura morfológica peculiar y adaptaciones locales, asociadas con la angulación del cuello y los músculos de la aleta dorsal, respectivamente. El tamaño vertebral disminuye desde elemento más anterior hacia atrás, con algunas variaciones locales en la zona dorsal. Los principales cambios están relacionados con la variación alométrica en la región neural. Los elementos caudales se caracterizan por una marcada disminución de tamaño, con los consiguientes cambios en la formas aloméricas de rotación de la abertura vertebral posterior. Esta trayectoria alométrica conduce a una flexión ventral natural de la cola, promoviendo su función prensil. Este estudio morfológico sugiere una interesante red estructural entre postura, locomoción y la anatomía vertebral junto a los principales cambios funcionales en la biomecánica de Hippocampus. La morfometría geométrica es bastante adecuada para los estudios con enfoques metaméricos, en términos de variación de serie y adaptaciones funcionales.

Skeletal Organization of Caudal Fin in Syngnathus abaster (Osteichthyes, Syngnathidae) / Organización Esqueletal de la Paleta Caudal en Syngnathus abaster (Osteichthyes, Syngnathidae)

The caudal skeleton is one of the most important tools to evaluate teleostean interrelationships, but just the anatomy of few species have been investigated. Black-striped pipefish (Syngnathus abaster) has never been examined for its caudal morphology. Here, red alizaryn is applied as specific staining for calcified structures, providing a better understanding of caudal skeleton organization. Our results and interpretation just partially fit with previous knowledge on other Syngnathus species. This emphasizes both the importance of facing same problems with new technics and methodologies, and the existence of a heterogeneous situation between close-related taxa.

El esqueleto caudal es una de las más importantes herramientas para evaluar las interrelaciones de teleósteros, pero la anatomía de pocas especies han sido investigada. Nunca se ha estudiado la morfología de la cauda del pez aguja de río de rayas negras (Syngnathus abaster). Se usó una técnica específica de tinción con alizarina roja, para visualizar estructuras calcificadas, obteniéndose un gran conocimiento de la organización del esqueleto caudal. Los resultados e interpretación de ellos coinciden parcialmente con los hallazgos en otras especies de Syngnathus. En ambos casos se enfatiza la importancia de tratar los problemas con nuevas técnicas y metodologías.