To date, most habitat models of cetaceans have relied on static and oceanographic covariates, and very few have related cetaceans directly to the distribution of their prey, as a result of the ...limited availability of prey data. By simulating the distribution of six functional micronekton groups between the surface and ≃1,000 m deep, the SEAPODYM model provides valuable insights into prey distributions. We used SEAPODYM outputs to investigate the habitat of three cetacean guilds with increasing energy requirements: sperm and beaked whales, Globicephalinae and Delphininae. We expected High Energy Requirements cetaceans to preferentially forage in habitats of high prey biomass and/or production, where they might easily meet their high energetic needs, and Low Energy Requirements cetaceans to forage in habitats of either high or low prey biomass and/or production. Cetacean sightings were collected from dedicated aerial surveys in the South West Indian Ocean (SWIO) and French Polynesia (FP). We examined cetacean densities in relation to simulated distributions of their potential prey using Generalised Additive Models and predicted their habitats in both regions. Results supported their known diving abilities, with Delphininae mostly related to prey present in the upper layers of the water column, and Globicephalinae and sperm and beaked whales also related to prey present in deeper layers. Explained deviances ranged from 9% for sperm and beaked whales in the SWIO to 47% for Globicephalinae in FP. Delphininae and Globicephalinae appeared to select areas where high prey biomass and/or production were available at shallow depths. In contrast, sperm and beaked whales showed less clear habitat selection. Using simulated prey distributions as predictors in cetacean habitat models is crucial to understand their strategies of habitat selection in the three dimensions of the ocean.
Tail flukes as well as the dorsal fin are the apomorphic traits of cetaceans which appeared during the evolutionary process of adaptation to the aquatic life. Both appendages present a wing-like ...shape associated with lift generation and low drag. We hypothesized that the evolution of fins as lifting structures led to a generic wing design, where the dimensionless parameters of the fin cross-sections are invariant with respect to the body length and taxonomy of small cetaceans (Hypothesis I). We also hypothesized that constraints on variability of a generic fin shape are associated with the primary function of the fin as a fixed or flapping hydrofoil (Hypothesis II). To verify these hypotheses, we examined how the variation in the fin’s morphological traits is linked to the primary function, species and body length. Hydrodynamic characteristics of the fin cross-sections were examined with the CFD software and compared with similar engineered airfoils. Generic wing design of both fins was found in a wing-like planform and a streamlined cross-sectional geometry optimized for lift generation. Divergence in a generic fin shape both on the planform and cross-sectional level was found to be related with the fin specialization in fixed or flapping hydrofoil function. Cross-sections of the dorsal fin were found to be optimized for the narrow range of small angles of attack. Cross-sections of tail flukes were found to be more stable for higher angles of attack and had gradual stall characteristics. The obtained results provide an insight into the divergent evolutionary pathways of a generic wing-like shape of the fins of cetaceans under specific demands of thrust production, swimming stability and turning control.
Despite large efforts, datasets with few sightings are often available for rare species of marine megafauna that typically live at low densities. This paucity of data makes modelling the habitat of ...these taxa particularly challenging. We tested the predictive performance of different types of species distribution models fitted to decreasing numbers of sightings. Generalised additive models (GAMs) with three different residual distributions and the presence only model MaxEnt were tested on two megafauna case studies differing in both the number of sightings and ecological niches. From a dolphin (277 sightings) and an auk (1,455 sightings) datasets, we simulated rarity with a sighting thinning protocol by random sampling (without replacement) of a decreasing fraction of sightings. Better prediction of the distribution of a rarely sighted species occupying a narrow habitat (auk dataset) was expected compared to the distribution of a rarely sighted species occupying a broad habitat (dolphin dataset). We used the original datasets to set up a baseline model and fitted additional models on fewer sightings but keeping effort constant. Model predictive performance was assessed with mean squared error and area under the curve. Predictions provided by the models fitted to the thinned-out datasets were better than a homogeneous spatial distribution down to a threshold of approximately 30 sightings for a GAM with a Tweedie distribution and approximately 130 sightings for the other models. Thinning the sighting data for the taxon with narrower habitats seemed to be less detrimental to model predictive performance than for the broader habitat taxon. To generate reliable habitat modelling predictions for rarely sighted marine predators, our results suggest (1) using GAMs with a Tweedie distribution with presence-absence data and (2) implementing, as a conservative empirical measure, at least 50 sightings in the models.
Understanding the mechanisms that drive prey selection is a major challenge in foraging ecology. Most studies of foraging strategies have focused on behavioural costs, and have generally failed to ...recognize that differences in the quality of prey may be as important to predators as the costs of acquisition. Here, we tested whether there is a relationship between the quality of diets (kJ · g(-1)) consumed by cetaceans in the North Atlantic and their metabolic costs of living as estimated by indicators of muscle performance (mitochondrial density, n = 60, and lipid content, n = 37). We found that the cost of living of 11 cetacean species is tightly coupled with the quality of prey they consume. This relationship between diet quality and cost of living appears to be independent of phylogeny and body size, and runs counter to predictions that stem from the well-known scaling relationships between mass and metabolic rates. Our finding suggests that the quality of prey rather than the sheer quantity of food is a major determinant of foraging strategies employed by predators to meet their specific energy requirements. This predator-specific dependence on food quality appears to reflect the evolution of ecological strategies at a species level, and has implications for risk assessment associated with the consequences of changing the quality and quantities of prey available to top predators in marine ecosystems.
The Mediterranean sperm whale population,
Physeter macrocephalus
, is listed as endangered due to population decline caused by human activities. To mitigate the impact of these activities, accurate ...knowledge of their distribution and abundance is crucial. During their long dives, sperm whales are not available to visual observation, but since they produce sounds when they dive, they are available to acoustic detection. Therefore, we aimed to use towed acoustic data to model their habitat and fill the knowledge gap on Mediterranean sperm whale distribution. Generalised additive models were used to link the number of sperm whales detected acoustically during the ACCOBAMS Survey Initiative in 2018 with different environmental variables integrated over different depth classes, encompassing the depth range used by the species for foraging. Sperm whale distribution was influenced by water temperature at the bottom, eddy kinetic energy between 200 and 600 m, as well as gradients of sea surface temperature and chlorophyll-a concentrations. The abundance of sperm whales was estimated at 2,959 individuals 2,077 - 4,265 in the sampled areas of the Mediterranean Sea. We predicted that sperm whales were mainly distributed in summer along the continental slope of the north-western Mediterranean basin from the Balearic Islands to the Ligurian Sea and off the Algerian coast. They were present throughout the western Mediterranean Sea and in the northern Ionian Sea. In contrast, predicted densities were low in the eastern part of the Mediterranean Sea. The use of acoustic data compensated for the main difficulty in studying sperm whales, the unavailability of animals at the surface during visual observation and the paucity of visual data. We thus encourage more systematic use of passive acoustics to study sperm whale distribution. The model highlighted a higher concentration of sperm whales in the western Mediterranean basin than in the eastern basin in summer, opening up avenues to improve the conservation of this endangered Mediterranean sub-population.
Spitz, J., Mourocq, E., Schoen, V., and Ridoux, V. 2010. Proximate composition and energy content of forage species from the Bay of Biscay: high- or low-quality food? – ICES Journal of Marine ...Science, 67: 909–915.Collapses of high-energy dense concentrations of prey species induce negative effects on populations of top predators. Knowledge of prey quality appears to be crucial in ecosystem modelling and management. The aim here was to provide baseline data of forage species quality in the Bay of Biscay. Proximate composition (water, ash, protein, and lipid) and energy content have been determined to assess the quality of 78 species, including jellyfish, crustaceans, cephalopods, cartilaginous fish, and bony fish. Results show broad variations between species, with energy densities ranging from 2 to 10 kJ g−1. Lipids are the most structuring component and largely determined prey quality, and prey species are not necessarily interchangeable for the fulfilment of a predator's energy and food requirements. In ecosystem models, therefore, multispecies compartments of forage organisms would ideally be constituted using prey species of equivalent quality and hence of equivalent benefit to top predators.
Individual competitiveness conditions access to resources when they are limited. Immature individuals that are less skilled than adults have to adapt their foraging strategies to survive. Among ...strategies to reduce competition, spatial segregation has been widely demonstrated. However, the use of spatial segregation by immatures to limit intra‐specific competition with adults has rarely been tested.
In this study, we investigated and compared habitat preferences and distributions of free‐ranging immature and breeding adult northern gannets Morus bassanus in order to determine whether they compete for similar habitats during the year, and if this results in a spatial segregation between birds of different age groups.
Based on > 66 000 km of aerial surveys conducted in the North‐East Atlantic Ocean during winter and summer 2012, habitats selected by immatures and adult birds were modelled independently, linking gannet density to a set of oceanographic and physiographic predictors. Their large‐scale seasonal distribution was then predicted.
We found that gannets displayed a strong season‐dependent competition between immatures and adults, as a consequence of immatures and adults using similar habitats in both summer and winter. During summer, when adults are constrained by reproduction, both groups were spatially highly segregated despite similar habitat preferences (thermal fronts), with youngest individuals selecting habitats out of range of central‐place foragers, highlighting intra‐specific competition. Contrastingly during winter, when reproductive constraints disappear, immature and adult distributions largely overlapped.
Our study provides new insights into the role played by age, foraging experience and reproductive constraints on the distribution of marine predators. More specifically, these results highlight in seabirds how the youngest fraction mitigates, through spatial segregation, the competition with experienced adults, and suggest a progressive strategy along the maturation process.
Estimating the detection probability of small cetaceans using either visual or acoustic surveys is difficult because they do not surface or vocalise continuously and can be imperceptible to an ...observer or hydrophone. Animals seen at the surface may have lower vocalisation rates, while submerged individuals may be more vocally active. This study aims to estimate visual, acoustic and combined detection probability by using Mark-Recapture Distance Sampling (MRDS) methodology. We used vessel-based visual sightings and acoustic data (based on click identification) collected simultaneously during the ACCOBAMS Survey Initiative in summer 2018 onboard the
R/V Song of the Whale
. This study focused on small cetaceans in the Mediterranean Sea, including the most commonly-encountered species, the striped dolphin
(Stenella coeruleoalba)
. We identified duplicate events between visual and acoustic platforms using a decision tree based on time and distance thresholds to estimate
g
(0) (the detection probability on the trackline) for small cetaceans. A total of 30 duplicate events were identified from 107 and 109 events identified by the visual and acoustic platforms respectively. We tested the models with two key functions. With a hazard-rate key function, the
g
(0) was estimated at 0.52 (CV=21.0%) for both platforms combined, 0.29 (CV=25.6%) for the visual platform and 0.32 (CV=25.1%) for the acoustic platform. With a half-normal key function,
g
(0) was estimated at 0.51 (CV = 21.7%) for both platforms combined, 0.29 (CV = 25.6%) for the visual platform and 0.33 (CV = 23.2%) for the acoustic platform. Our results illustrate that passive acoustic monitoring can be used as an independent platform in MRDS to estimate the detection probability. Our estimate of
g
(0) was well below 1, far from the perfect detection commonly assumed for abundance estimation. Without correction for detection biases, total abundance would be underestimated by a factor of two when using both acoustic and visual data. This highlights the importance of using dual-platform surveys to estimate detection probability in order to improve abundance estimates and conservation efforts.
Top predators need to develop optimal strategies of resources and habitats utilization in order to optimize their foraging success. At the individual scale, a predator has to maximize his intake of ...food while minimizing his cost of foraging to optimize his energetic gain. At the ecosystem scale, we hypothesized that foraging strategies of predators also respond to their general energetic constraints. Predators with energetically costly lifestyles may be constrained to select high quality habitats whereas more phlegmatic predators may occupy both low and high quality habitats. The objectives of this study were 1) to investigate predator responses to heterogeneity in habitat quality with reference to their energetic strategies and 2) to evaluate their responses to contemporaneous versus averaged habitat quality. We collected cetacean and seabird data from an aerial survey in the Southwest Indian Ocean, a region characterized by heterogeneous oceanographic conditions. We classified cetaceans and seabirds into energetic guilds and described their habitats using remotely sensed covariates at contemporaneous and time-averaged resolutions and static covariates. We used generalized additive models to predict their habitats at the regional scale. Strategies of habitat utilization appeared in accordance with predators energetic constraints. Cetaceans responded to the heterogeneity in habitat quality, with higher densities predicted in more productive areas. However, the costly Delphininae appeared to be more dependent on habitat quality (showing a 1-to-13 ratio between the lowest and highest density sectors) than the more phlegmatic sperm and beaked whales (showing only a 1-to-3 ratio). For seabirds, predictions primarily reflected colony locations, although the colony effect was stronger for costly seabirds. Moreover, our results suggest that predators may respond better to persistent oceanographic features. To provide a third dimension to habitat quality, cetacean strategies of utilization of the vertical habitat could be related to the distribution of micronekton in the water column.
Overfishing and ocean warming are drastically altering the community composition and size structure of marine ecosystems, eliminating large bodied species 1. Against a backdrop of such environmental ...change, the heaviest of all bony fish, the ocean sunfish (Mola mola), seems an improbable survivor. Indeed this indolent giant is killed globally as bycatch, and is listed as ‘Vulnerable’2. We undertook the most extensive aerial surveys of sunfish ever conducted and found surprisingly high abundances off the Atlantic and Mediterranean coasts of Western Europe. With up to 475 individuals per 100 km2, these figures are one order of magnitude higher than abundance estimates for other areas 3–5. Using bioenergetic modelling, we estimate that each sunfish requires 71 kg day–1 of jellyfish, a biomass intake more than an order of magnitude greater than predicted for a similarly sized teleost. Scaled up to the population level, this equates to a remarkable 20,774 tonnes day–1 of predated jellyfish across our study area in summer. Sunfish abundance may be facilitated by overfishing and ocean warming, which together cause reduced predation of sunfish by sharks and elevated jellyfish biomass. Our combined survey and bioenergetic data provide the first-ever estimate of spatialized ocean sunfish daily food requirements, and stress the importance of this species as a global indicator for the ‘rise of slime’. This hypothesis posits that, in an overfished world ocean exposed to global warming, gelatinous zooplankton should flourish, to the detriment of other mesotrophic species such as small pelagic fish, causing irreversible trophic cascades as well as a series of other environmental and economic issues.
The Ocean Sunfish is the heaviest of all bony fish, and is listed as vulnerable by the IUCN. Grémillet et al. conducted aerial surveys off the Atlantic and Mediterranean coasts of Western Europe, finding up to 475 sunfish per 100 km2. The sunfish population in this region is estimated to consume over 20,000 tonnes of jellyfish per day in summer.
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