Diving behaviour of female northern rockhopper penguins, Eudyptes chrysocome moseleyi, during the brooding period at Amsterdam Island (Southern Indian Ocean)

The pattern and characteristics of diving in 14 female northern rockhopper penguins, Eudyptes chrysocome moseleyi, were studied at Amsterdam Island (37°50′S; 77°31′E) during the guard stage, using electronic time–depth recorders. Twenty-nine foraging trips (27 daily foraging trips and two longer trips including one night) with a total of 16 572 dives of ≥3 m were recorded. Females typically left the colony at dawn and returned in the late afternoon, spending an average of 12 h at sea, during which they performed ∼550 dives. They were essentially inshore foragers (mean estimated foraging range 6 km), and mainly preyed upon the pelagic euphausiid Thysanoessa gregaria, fishes and squid being only minor components of the diet. Mean dive depth, dive duration, and post-dive intervals were 18.4 m (max. depth 109 m), 57 s (max. dive duration 168 s), and 21 s (37% of dive duration), respectively. Descent and ascent rates averaged 1.2 and 1.0 ms⁻¹ and were, together with dive duration, significantly correlated with dive depth. Birds spent 18% of their total diving time in dives reaching 15 to 20 m, and the mean maximum diving efficiency (bottom time:dive cycle duration) occurred for dives reaching 15 to 35 m. The most remarkable feature of diving behaviour in northern rockhopper penguins was the high percentage of time spent diving during daily foraging trips (on average, 69% of their time at sea); this was mainly due to a high dive frequency (∼44 dives per hour), which explained the high total vertical distance travelled during one trip (18 km on average). Diving activity at night was greatly reduced, suggesting that, as other penguins, E. chrysocome moseleyi are essentially diurnal, and locate prey using visual cues. Received: 9 December 1998 / Accepted: 3 March 1999     

Diving behaviour of gentoo penguins,Pygoscelis papua; factors keeping dive profiles in shape

The foraging ecology of seven Gentoo penguins,Pygoscelis papua, breeding at Ardley Island, Antarctica was studied using animal-attached devices which recorded swimming speed, heading and dive depth. Reconstruction of the foraging routes by vectorial analysis of the data indicated that at no time did the birds forage on the sea bed. Swimming speed was relatively constant at 1.7 m s^-1, but rates of descent and ascent in the water column during dives increased with increasing maximum dive depth due to changes in descent and ascent angles. The amount of time spent discending and ascending in the water column increased with maximum dive depth as did the duration spent at the point of maximum depth. Dive profiles were essentially either U-shaped (flat-bottomed dives), or V-shaped (bounce dives). Development of a model based on simple probability theory indicated that the optimal dive profile to maximize the chances of prey acquisition depends on vertical prey distribution and on the visual capabilities of the birds with respect to descent and ascent angles.     

Foraging behaviour and reproductive success in Magellanic penguins (Spheniscus magellanicus): a comparative study of two colonies in southern Chile

Marine Biology - During the breeding season 1996/97 we compared the foraging and diving behaviour of adult Magellanic penguins (Spheniscus magellanicus), growth rates of their chicks and their...     

Seasonal changes in the diving parameters of king penguins (Aptenodytes patagonicus)

Contrasting conditions at-sea are likely to affect the foraging behaviour of seabirds. However, the effect of season on the dive parameters of penguins is poorly known. We report here on an extensive study of the diving behaviour of king penguins (Aptenodytes patagonicus) over the bird's complete annual cycle at the Crozet Islands. Time-depth recorders were used to record dive duration, bottom duration, post-dive interval, ascent rate and descent rate in breeding adults during different seasons in 1995 and 1996. Seasons included summer (n=6, incubation; n=6, chick brooding), autumn and winter (n=5 and n=3, respectively, chick at the crèche stage), and spring (n=4, birds at the post-moult stage). In all seasons dive duration increased with dive depth, but, for a given depth, dives were longer in winter (6.8 min when averaged over the 100-210 m depth layer) than in spring (4.6 min) and summer (4.4 min). The time spent at the bottom of the dives, which probably represents a substantial part of the feeding time, was much longer in winter (2.5 min per dive for dives over the 100-210 m layer) than during other seasons (1.0-1.4 min), i.e. there was a 2.5-fold augmentation for similar diving depths. Ascent and descent rates increased with increasing dive depth, but no difference in the relationships between rates of ascent and descent and dive depth was found among seasons. Furthermore, for all dive depths, ascent and descent rates were independent of the bottom duration. In all seasons post-dive intervals increased with dive duration and with dive depth, but they were longer in spring (2.3 min for dives over the 100-210 m layer) and summer than in autumn and winter (1.6-1.8 min). The diving efficiency decreased with increasing dive depth and was higher in autumn and winter (0.22-0.29) than in summer and spring (0.15-0.18). The large increase in bottom and dive duration from spring to winter is in agreement with the seasonal drop in prey density, with penguins spending more time searching for prey. In contrast, the consistency of the vertical velocity during contrasting conditions at-sea suggests that the transit time to depth is an important component of the foraging behaviour (scanning of the water column) that is independent of the prey availability. The time budget of the penguins during diving in a fluctuating environment appears to vary primarily during the bottom phase of the dives, with bottom duration increasing with diminishing prey supplies, while post-dive intervals shorten in the same time.     

Satellite tracking of Humboldt penguins (Spheniscus humboldti) in northern Chile

During the El Niño of 1982/1983, the Humboldt penguin population diminished dramatically in the whole distributional area of the species. Recovery of the population was slow since 1983 and it has been suggested that large numbers of Humboldt penguins die at sea, entangled in nets, or starve to death, even during non-“El Niño” years. We were able to determine for the first time, how Humboldt penguins on Pan de Azúcar Island (26°S; 72°W) utilize their marine habitat and where their feeding areas lie. For this purpose we employed two streamlined Argos satellite transmitters during the 1994/1995 and 1995/1996 breeding seasons, respectively. Mean travelling speed of Humboldt penguins during foraging trips was 0.94?m s^?1 and 50% of bird positions were located within 5?km of the island (90% within 35?km). Total area covered by Humboldt penguins foraging from Pan de Azúcar Island was 12?255?km². Satellite transmitters also recorded dive duration; penguins spent on average 7.8 to 9?h diving per foraging day but showed no preferences for particular feeding areas. Mean daily dive durations (4-d mean) recorded during the 1994/1995 breeding season were positively correlated between birds. Significant correlation between dive duration and sea surface temperature anomalies and negative correlation between dive duration and fishery landings at nearby Caldera harbour indicate that the 1994/1995 increase in foraging effort was a response to deteriorating prey availability. Sea surface temperatures during the 1995/1996 breeding season were colder than average, and we observed no trends in bird diving activities.     

The section Digitatae of the genus Laminaria (Phaeophyta) in the northern and southern Atlantic: crossing experiments and temperature responses

Hybridization experiments between seven north and south Atlantic Laminaria species were carried out. Morphologically normal F₁ sporophytes developed from the following crosses among south Atlantic species: L. pallida x L. schinzii, L. pallida x L. abyssalis and L. schinzii x L. abyssalis. Normal F₁ sporophytes also resulted from the crosses L. digitata (north Atlantic) x L. pallida (south Atlantic) and L. digitata (north Atlantic) x L. abyssalis (south Atlantic). Hybrids between north Atlantic L. ochroleuca and south Atlantic L. pallida, L. schinzii and L. abyssalis and between north Atlantic L. digitata and south Atlantic L. schinzii initially developed as normal sporophytes but became deformed later on and further development was retarded. No hybrids resulted from attempted crosses between northeastern Atlantic L. saccharina and L. abyssalis from Brazil. Temperature tolerance, relative growth rates and temperature demands for gametogenesis revealed the existence of a warm temperate group within the digitate Laminaria species consisting of L. ochroleuca, L. pallida, L. schinzii and L. abyssalis. Hybridization experiments and temperature responses suggest that north Atlantic L. digitata and L. ochroleuca are still similar to south Atlantic Laminaria species, confirming the speculation that a transequatorial migration of a warm-temperate L. ochroleuca-like ancestor may have taken place.     

Winter migration of magellanic penguins (Spheniscus magellanicus) from the southernmost distributional range

A total of 22 magellanic penguins (Spheniscus magellanicus) from Isla Martillo in the Beagle Channel, Argentina, were successfully satellite tracked in 2004 (n = 7), 2005 (n = 7) and 2006 (n = 8) to monitor their winter migration after moult. Only one magellanic penguin migrated northwards into the Pacific Ocean, whereas all others remained in the Atlantic Ocean. In general, these birds left the island in an easterly direction, rounded Cabo San Diego, the southeasterly tip of South America, and continued northwards occupying inshore waters mostly less than 50 km from the coast, only occasionally venturing further offshore. By the end of the transmission period, birds were still travelling northwards and the most northerly positions were obtained from birds located in the area of Peninsula Valdés, Argentina, at a latitude of around 42°S, some 1,500 km from their breeding site on Isla Martillo. The mean maximum distance to the breeding site was, however, only 624 ± 460 km. The mean minimum distance covered during the study period was 1,440 ± 685 km, which corresponded to a mean distance of 23.2 ± 6.6 km covered per day. The northbound migration of the penguins could be separated into periods of rapid movement, interspersed with periods during which the birds remained for some time in particular coastal regions. Areas with a high density of daily penguin positions were observed in three distinct areas: at the northeastern coast of Tierra del Fuego, at the southern entrance of Golfo San Jorge and to the northeast of the Peninsula Valdez. The observed migration pattern is presumably driven by the formation and subsequent dispersal of areas of enhanced productivity as the season progressed. Our findings also suggest that magellanic penguins are increasingly threatened by human activities in coastal areas as penguins migrate northwards.     

Effects of attached data-logging devices on little penguins (Eudyptula minor)

Data-logging devices are commonly used to study the foraging behaviour of individual seabirds. Such studies need to examine the potential effects of using devices on instrumented individuals, not only for ethical reasons but also to ensure the validity of data gathered. We studied the effects of two types of device (time-depth recorder and global positioning system) on little penguins (Eudyptula minor) during the 2010 and 2011 breeding season at Oamaru, New Zealand. Mixed-effect models were used to test for effects of devices by comparing changes in body weight, chick growth and breeding performance between instrumented and control individuals. We found no detectable effects of the attached devices on body weight change, hatching success, fledging success, chick growth parameters or adult survival. We conclude that it is possible to attach data-logging devices to adult little penguins for extended periods during the breeding season with minimal impacts.     

Behavioural switching in a central place forager: patterns of diving behaviour in the macaroni penguin (Eudyptes chrysolophus)

Recording the activity of animals as they migrate or forage has proven hugely advantageous to understanding how animals use their environment. Where animals cannot be directly observed, the problem remains of how to identify distinct behaviours that represent an animal’s decision-making process. An excellent example of this problem is that of foraging penguins, which travel to sea to find prey to provision their young. Without direct sampling of the prey field, we cannot calibrate patterns of movement with prey capture, and therefore we cannot determine how different activities link to decision-making. To overcome this, we use a hidden markov model (HMM), which is a machine-learning technique that seeks to identify the underlying states of a system from observable outputs. We apply HMM to determine classes of behaviour from repetitive dives. We take dive data from 103 breeding macaroni penguins at Bird Island, South Georgia, for which we have measures of weight gain over a trip. We identify two classes of behaviour; those of short-shallow and long-deep dives. Using these two behaviours, we calculate the transition probabilities between these states and analyse these data to determine what predicts variation in the transition probabilities. We found that the stage of reproduction during a season, the sex and year of an individual influenced the probability of transition between long-deep and short-shallow sequential dives. We also found differences in the hourly transition rates between the four reproductive stages (incubation, broodguard, crèche and premoult) over a daily cycle. We conclude that this application of HMMs for behavioural switching is potentially useful for other species and other types of recorded behaviour.     

Diving behavior of immature hawksbills (Eretmochelys imbricata) in a Caribbean cliff-wall habitat

Time-depth recorders were attached to immature hawksbills (Eretmochelys imbricata Linnaeus, 1766) residing at the northwestern cliffs of Mona Island, Puerto Rico. Data on diving profiles obtained from four turtles of 32.0 to 53.4 cm straight carapace length revealed strong similarities in behavioral patterns. During daylight hours, turtles were active an average 8.4 h per day, surfacing 3.6% of the time. Length of foraging dives correlated with turtle size, with mean durations ranging from 8.6 to 14.0 min. Foraging dives, with a mean depth of 4.7 m, were associated with feeding on encrusting sponges. At night, turtles were mostly inactive, surfacing 1.8% of the time and with individual mean submergence intervals of between 30.4 and 37.1 min. From the surfacing behavior of turtles making deep and long dives, it is inferred that dives were aerobic, with the turtles making use of oxygen stores in addition to that of the lung.     

Trace metals in mussels (Mytilus edulis) from southwest Iceland

The temperate population of the tropical anemonefish Amphiprion clarkii has a seasonality of reproduction, larval settlement and growth. There was a considerable difference in size (15–47 mm) among 0-year olds in December, after the season of larval settlement and growth. This difference could be attributed mainly to a difference in the duration of the first growing season spent after settlement and a social inhibition of growth of the subordinate late settlers by frequent attacks by the dominant early settlers. The growth equation which was used to describe the growth of tagged individuals estimated that an anemonefish would get an opportunity to spawn within four of five years after settlement. Breeding adult pairs occupied almost all sea anemones essential to breeding within a 50×50 m study area, and all 17 new adults that matured from non-breeding juveniles to breeding adults between June 1983 and August 1985 were found in the territories from which either or both of the mated adults had disappeared. The new adults were large juveniles who had resided near those vacant territories before. These results suggest that the onset of breeding by an individual is not only determined by his age, but also by his ranking in the dominance hierarchy. Therefore, the difference in size among 0-year olds in December might give rise to the difference in age of the earliest breeding and might be a factor in making a female apply more reproductive effort at the beginning of the season.     

Tracking adult North Atlantic bluefin tuna (Thunnus thynnus) in the northwestern Atlantic using ultrasonic telemetry

Ultrasonic, depth-sensitive transmitters were used to track the horizontal and vertical movements, for up to 48 h, of 11 adult (136 to 340 kg estimated body mass) North Atlantic bluefin tuna (Thunnus thynnus Linnaeus). Fish were tracked in October 1995, September and October 1996, and August and September 1997 in the Gulf of Maine, northwestern Atlantic. The objective was to document the behavior of these fish and their schools in order to provide the spatial, temporal, and environmental information required for direct (i.e. fishery-independent) assessment of adult bluefin tuna abundance using aerial surveys. Transmitters were attached to free-swimming fish using a harpoon attachment technique, and all fish remained within the Gulf of Maine while being followed. Most of the bluefin tuna tagged on Stellwagen Bank or in Cape Cod Bay (and followed for at least 30 h) held a predominately easterly course with net horizontal displacements of up to 76 km d⁻¹. Mean (±SD) swimming depth for all fish was 14 ± 4.7 m and maximum depth for individuals ranged from 22 to 215 m. All but one fish made their deepest excursions, often single descents, at dawn and dusk. In general, adult bluefin tuna spent <8% of their time at the surface (0 to 1 m), <19% in the top 4 m, but >90% in the uppermost 30 m. Mean (±SD) speed over ground was 5.9 km h⁻¹, but for brief periods surpassed 20 to 31 km h⁻¹. Sea surface temperatures during tracking were 11.5 to 22.0 °C, and minimum temperatures encountered by the fish ranged from 6.0 to 9.0 °C. Tagged bluefin tuna and their schools frequented ocean fronts marked by mixed vertebrate feeding assemblages, which included sea birds, baleen whales, basking sharks, and other bluefin schools. Received: 19 July 1999 / Accepted: 25 March 2000     

Proximity to competitors changes secondary metabolites of non-indigenous cup corals, Tubastraea spp., in the southwest Atlantic

Competition for space changes species’ distributions and community organization on tropical rocky shores, and the presence of secondary metabolites in the tissues of non-indigenous species may aid them in establishing and expanding their range through negative competitive interactions. The aim of this study was to describe the range of chemical substances produced by the non-indigenous cup corals Tubastraea coccinea and T. tagusensis and to test whether they varied in the field when the corals were placed in proximity to two local competitors. Cholest-5-en-3β-ol and 9-octadecanoic acid were two common secondary metabolites found in the tissues of Tubastraea. In the competition interaction experiment, necrosis was detected on the tissues of the coral Mussismilia hispida, and this species induced variation in sterol, alkaloid, and fatty acid production in Tubastraea tissues. In contrast, a sponge overgrew Tubastraea colonies. These results indicate that chemical defense may contribute to the ability of these non-indigenous corals to invade native communities.     

Population differentiation of the Atlantic spotted dolphin (Stenella frontalis) in the western North Atlantic, including the Gulf of Mexico

Information about the genetic population structure of the Atlantic spotted dolphin [Stenella frontalis (G. Cuvier 1829)] in the western North Atlantic would greatly improve conservation and management of this species in USA waters. To this end, mitochondrial control region sequences and five nuclear microsatellite loci were used to test for genetic differentiation of Atlantic spotted dolphins in the western North Atlantic, including the Gulf of Mexico (n=199). Skin tissue samples were collected from 1994–2000. Significant heterozygote deficiencies in three microsatellite loci within samples collected off the eastern USA coast prompted investigation of a possible Wahlund effect, resulting in evidence for previously unsuspected population subdivision in this region. In subsequent analyses including three putative populations, two in the western North Atlantic (n=38, n=85) and one in the Gulf of Mexico (n=76), significant genetic differentiation was detected for both nuclear DNA (R _ST=0.096, P≤0.0001) and mitochondrial DNA (Φ _ST=0.215, P≤0.0001), as well as for all pair-wise population comparisons for both markers. This genetic evidence for population differentiation coupled to known biogeographic transition zones at Cape Hatteras, North Carolina and Cape Canaveral, Florida, USA, evidence of female philopatry, and preliminary support for significant genetic differences between previously documented morphotypes of Atlantic spotted dolphins in coastal and offshore waters all indicate that the biology and life history of this species is more complex than previously assumed. Assumptions of large, panmictic populations might not be accurate in other areas where S. frontalis is continuously distributed (e.g., eastern Atlantic), and could have a detrimental effect on long-term viability and maintenance of genetic diversity in this species in regions where incidental human-induced mortality occurs.

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Homing of eels (Anguilla anguilla) in the southern North Sea

Tagging and transplantation experiments on a total of 1538 cels (Anguilla anguilla L.) caught in the German Bight near the islands Helgoland and Föhr, as well as in the area northwest of the Elbe estuary near Cuxhaven and in the port of Hamburg, resulted in recapture of 77 eels (5.0%). Of the recaptured eels 64% were reported from the area of their original capture. Some homed from distances of 180 km. Eels tagged and released within the area of original capture (control experiments) were recaptured only in these areas. A maximum migration speed of at least 1 km/h was found in one case, and of 3 km/h in another. Average speed of migration may, however, have been much lower, since in most other cases recaptures were reported at least 2 weeks after release. The homing abilities demonstrated cannot in all cases be accounted for by olfactory sensitivity alone; home areas were in some cases located in such a way that the water circulation in the southern North Sea could not have provided any lead for the migrating eels.     

Population genetics of the nurse shark (Ginglymostoma cirratum) in the western Atlantic

The nurse shark, Ginglymostoma cirratum, inhabits shallow, tropical, and subtropical waters in the Atlantic and the eastern Pacific. Unlike many other species of sharks, nurse sharks are remarkably sedentary. We assayed the mitochondrial control region and eight microsatellite loci from individuals collected primarily in the western Atlantic to estimate the degree of population subdivision. Two individuals from the eastern Atlantic and one from the Pacific coast of Panama also were genotyped. Overall, the mtDNA haplotype (h = 48 ± 5%) and nucleotide (π = 0.08 ± 0.06%) diversities were low. The microsatellite data mirror the mitochondrial results with the average number of alleles ([`(N)]<sub>A</sub> \bar{N}_{A}  = 9) and observed heterozygosity ([`(H)]_O \bar{H}_{O}  = 0.58) both low. The low levels of diversity seen in both the mtDNA and the microsatellite may be due to historical sea level fluctuations and concomitant loss of shallow water habitat. Eight of the 10 pair-wise western Atlantic F _ST estimates for mtDNA indicated significant genetic subdivision. Pair-wise F _ST values for the microsatellite loci indicated a similar pattern as the mtDNA. The western Atlantic population of nurse sharks is genetically subdivided with the strongest separation seen between the offshore islands and mainland Brazil, likely due to deep water acting as a barrier to dispersal. The eastern and western Atlantic populations were closely related. The eastern Pacific individual is quite different from Atlantic individuals and may be a cryptic, sister species.     

西南岩溶山区景观生态特征与景观生态建设

探讨了西南岩溶山区景观生态结构与功能以及景观生态过程的不同于其他区域的基本特征，尝试从景观角度认识西南岩溶区生态空间特性与演化过程。受地质地貌和土地利用的影响。西南岩溶山区景观基质脆弱、景观格局分异明显，具破碎化与景观结构粗粒化、景观动态变化范围大且速度快、整体退化明显的特点，以石漠化为特征的景观演化削弱了岩溶生态系统的生态服务功能。岩溶山区的景观生态建设须充分认识岩溶山区资源环境优势，充分利用当地的适生植物资源，协调生态效应和水文效应，促进水土资源的有效保持向高效利用发展，形成新的高效、和谐的人工自然景观。今后应把景观格局与生态过程结合起来、与人文因子和地学背景结合起来，加强小尺度的景观变化研究。     

Sardine (Sardina pilchardus) spawning seasonality in European waters of the northeast Atlantic

Egg data from ichthyoplankton monitoring sites in the western English Channel (1988–2003) and northern Spain (1990–2000) and macroscopic maturity data from biological samples of purse seine landings in western and southern Iberia (1980–2004) are used to describe the spawning seasonality of sardine (Sardina pilchardus) in European waters of the northeast Atlantic using generalised additive models. The fitted models reveal a double peak in spawning activity during early summer and autumn in the western Channel, a wider spring peak off northern Spain and a broad winter season in the western and southern Iberian Peninsula. At all sites, a high probability of spawning activity was observed over at least 3 months of the year, with the duration of the season increasing with both decreasing latitude and increasing fish size. Off western and southern Iberia there are indications that the spawning season has been of longer duration in recent years for all size classes (reaching in some cases 8 months of the year for large fish). These patterns are in general agreement with existing literature and theoretical expectations of sardine spawning being driven locally by the seasonal cycle of water temperature, assuming preferences for spawning at 14 –15°C and avoidance for temperatures below 12°C and above 16°C. Regional quotient plots indicated that spawning tolerance to higher temperatures increases progressively with decreasing latitude. Despite the weak evidence for geographical differences in temperature tolerance that may have some genetic origin, the degree of spatio-temporal overlap in sardine-spawning activity within Atlantic European waters is unlikely to promote any reproductive isolation in that area.     

Distribution of nutrients and AOU in the Mediterranean Water (North Atlantic Ocean)

Water samples were collected in the Northeast Atlantic Ocean (33-37^° N and 11-16^° W) in May 1983, and vertical profiles of chemical and physical parameters allowed us to identify the different water masses. A significant difference in salinity and its maximum can be found at different places and depths, as a consequence of the spreading of the Mediterranean Water (MW) from the Strait of Gibraltar. The apparent oxygen utilization shows a correlation with nutrients in North Atlantic Central Water. The relative percentages of water masses were assessed in order to examine the depth of maximum influence of MW as a function of latitude and to apply this knowledge to the black scabbard fish long-line strategy. The long-line fishing depth location was adapted to the specific MW distribution of the different banks with improvements in the mean catch efficiency. The thickness of the MW prevalence layer has a pronounced decrease to the south. MW influence clearly sinks and decrease to south-west. The highest percentage of the MW was found in the Gorringe Bank area and the lowest in the Madeira area.     

Antarctic krill aggregation characteristics from acoustic observations in the Southwest Atlantic Ocean

The distributional features and physical characteristics of 4830 krill (Euphausia superba Dana) aggregations detected acoustically in the Southwest Atlantic between 26 January and 21 February 1981 are described. Results are compared with aggregations detected in the Indian Ocean. Aggregations in the Atlantic were larger, closer to the surface and to each other than in the Indian Ocean. Similar patterns in the distribution of aggregation spacing along survey transects were found in the two areas, although the pattern of spacings in the Atlantic indicates differences in the scale of aggregation. Serial interdependence of aggregation variables was minimal in the Atlantic, with aggregation thickness, length and spacing showing weak inter-relationships. Weak functional association, between water depth and aggregation thickness was evident. Investigation of variability in aggregation structure in relation to prevailing environmental conditions gave equivocal results and no clear association between any aggregation variable and prevailing hydrography was observed. The implications of these results for future studies on krill aggregation are discussed in relation to a conceptual framework which was developed from the present results and aimed at linking krill aggregation characteristics to environmental features.