Movements of whale sharks (Rhincodon typus) tagged at Ningaloo Reef, Western Australia

Whale sharks (Rhincodon typus Smith) aggregate seasonally (March–June) to feed in coastal waters off Ningaloo Reef, Western Australia. Pop-up archival tags were attached to 19 individuals (total lengths 4.5–11.0 m) at this location in early May of 2003 and 2004 to examine their horizontal and vertical movements. The long-term movement patterns of six whale sharks were documented, all of which travelled northeast into the Indian Ocean after departing Ningaloo Reef. They used both inshore and offshore habitats and made extensive vertical movements, occasionally to a depth of at least 980 m. Frequent up-and-down movements, diel vertical migration, and crepuscular descents were evident in the depth records. The sharks experienced ambient temperatures ranging between 4.2 and 28.7°C and encountered gradients of up to 20.8°C on dives.     

Diel vertical movements of adult male dolphinfish (Coryphaena hippurus) in the western central Atlantic as determined by use of pop-up satellite archival transmitters

The vertical movements of six adult male dolphinfish (Coryphaena hippurus) (95–120 cm estimated fork length), caught using standard sportfishing methods, were investigated using high-rate single-point pop-up satellite archival transmitters from 2005 to 2011 in the western central Atlantic. Data revealed a diel activity pattern within the mixed surface layer with dives below the thermocline suggesting temperature is not a barrier to vertical movements for short periods of time. Dolphinfish were tracked for periods of 4.96–30.24 day (Σ = 83.37 day), reaching depths >200 m, and in temperatures ranging from 16.20 to 30.87 °C. The six tags allowed comprehensive vertical movement analyses by time of day, duration at depth, and based on vertical movement patterns. The longest (>60 min), deepest (>30 m), and most extensive vertical movement patterns occurred during night rather than day, with the most time spent near the surface during the day. Dolphinfish spent 66 % of their time in the surface layer (0–9.9 m) and only one individual spent 8 % of the monitoring period diving >8 °C from the maximum surface temperatures recorded while tracked. Two tags were analyzed based on lunar phase and revealed contrasting relationships between vertical movements during new and full phases. Our results suggest dolphinfish vertically shift between surface and at-depth feeding strategies to exploit aggregating epipelagic and mesopelagic prey items leading to predictable diel vertical movements.     

The effects of shark cage-diving operations on the behaviour and movements of white sharks, Carcharodon carcharias, at the Neptune Islands, South Australia

The attraction or provisioning of sharks for the purpose of tourism is a lucrative and popular industry that remains controversial regarding its possible risks to target species and impacts on local ecosystems. The long-term impacts of such activities on the behaviour and movement patterns of sharks have typically been difficult to establish as most studies investigate contemporary behaviour concurrent with existing operations and thus have no comparative base from which to compare effects. We compared patterns of residency and behaviour of acoustic-tagged white sharks at the Neptune Islands in South Australia between periods before and after an abrupt and sustained doubling of cage-diving effort that occurred in 2007. The number of sharks reported by cage-dive operators significantly increased after 2007. Comparisons also revealed there were significant increases in sharks’ periods of residency, the periods spent within areas where shark cage-diving operations occur and changes in sharks’ diel pattern of habitat use. Changes were site-specific with no significant differences in shark behaviour revealed over the same period at an island group 12 km from regular shark cage-dive sites. The results suggest that cage-diving operations can lead to long-term changes in the site-specific behaviour of a highly vagile shark species which may need to be considered in the context of their conservation and in managing the impacts of the industry.     

Observations on the short-term movements and behaviour of whale sharks (Rhincodon typus) at Ningaloo Reef, Western Australia

The short-term movements and behaviour of whale sharks (Rhincodon typus Smith, 1828) during March 1994 and April 1997 are reported from data collected by acoustic tracking and archival tags at Ningaloo Reef on the north west coast of Western Australia. Sharks were tracked for up to 26 h and generally swam slowly at ≃0.7 m s⁻¹ parallel to the reef edge; occasionally they swam in a wide arc adjacent to passes in the reef. All tracked sharks made regular dives through the water column, mostly from the surface to near the bottom. These dives did not appear to be related to hydrographic features, and the sharks were probably searching the water column for food. Most sharks were accompanied by other fishes, usually the golden trevally Gnathanodon speciosus. Received: 19 January 1999 / Accepted: 22 June 1999     

Spatial assessment of fin whale hotspots and their association with krill within an important Antarctic feeding and fishing ground

Fin whale (Balaenoptera physalus quoyi) habitat use and its relationship to environmental conditions are generally unknown in the Southern Ocean, presenting challenges for predicting their seasonal occurrence and potential effects of fishing pressure and climate change on this endangered species. Using biological data collected during 14 shipboard surveys off the northern Antarctic Peninsula and oceanographic data from satellite remote sensing, we mapped the distribution of fin whale hotspots, Antarctic krill abundance (biomass from acoustics, concentrations from nets) and ocean conditions during mid- and late-summer to investigate the environmental determinants of whale hotspots. Generalized additive models (GAM) were used to test the hypothesis that intra-seasonal changes in fin whale hotspot distribution relate to sea surface temperature (SST), krill abundance and eddy kinetic energy (EKE). More whale hotspots (sightings and individuals) are observed during late- than mid-summer surveys. During mid-summer, hotspots occurred near Elephant Island while in late-summer they were distributed throughout the slope region in proximity to the mean location of the southern Antarctic Circumpolar Current Front. The spatial mean of EKE did not differ between mid- and late-summer surveys, but the spatial mean of SST was significantly warmer during late-summer. The GAM for mid-summer indicates that fin whale hotspots were positively related to SST, EKE and acoustically determined krill biomass. The GAM for late-summer indicates the hotspots were negatively related to net-based krill abundance and positively related to acoustic krill biomass and EKE. This study is important because environmental determinants of fin whale hotspots may be used as reference points for implementing future conservation plans for their recovering populations.     

Seasonal changes in movements and habitat preferences of the scalloped hammerhead shark (Sphyrna lewini) while refuging near an oceanic island

Movements and habitat preferences of sharks relative to a central location are widely documented for many species; however, the reasons for such behaviors are currently unknown. Do movements vary spatially or temporally or between individuals? Do sharks have seasonal habitat and environmental preferences or simply perform movements at random at any time of the year? To help understand requirements for the designation of critical habitats for an endangered top predator and to develop zoning and management plans for key habitats, we examined vertical and horizontal movements, and determined habitat and environmental preferences of scalloped hammerhead sharks (Sphyrna lewini). We tracked seven hammerheads for 19–96 h at Wolf Island (1.38ºN, 91.82ºW) between 2007 and 2009 using ultrasonic transmitters with depth and temperature sensors, and we profiled temperature through the water column. Movements of individual hammerheads fell in two classes: constrained (remaining near the island) and dispersive (moving offshore to pelagic environments). The central activity space or kernel off the southeast side of Wolf Island was small and common to most, but the area varied among individuals (mean ± SE 0.25 ± 0.2 km²), not exceeding 0.6 km² for any of the sharks, and not changing significantly between seasons. In general, hammerheads showed preference for the up-current habitat on the eastern side of Wolf Island in both the warm and cold seasons. However, the depth of sharks varied with season, apparently in response to seasonal changes in the vertical structure of temperature. Hammerheads performed frequent vertical excursions above the thermocline during offshore movements and, in general, were observed to prefer temperatures of 23–26 °C found above the thermocline. At times, though individuals moved into the thermocline and made brief dives below it. Our results provided evidence that hammerheads (1) are highly selective of location (i.e., habitat on up-current side of island) and depth (i.e., top of the thermocline) while refuging, where they may carry out essential activities such as cleaning and thermoregulation, and (2) perform exploratory vertical movements by diving the width of the mixed layer and occasionally diving below the thermocline while moving offshore, most likely for foraging.     

Geographical variation in the behaviour of a central place forager: Antarctic fur seals foraging in contrasting environments

Foragers show adaptive responses to changes within their environment, and such behavioural plasticity can be a significant driving force in speciation. We investigated how lactating Antarctic fur seals, Arctocephalus gazella, adapt their foraging within two contrasting ecosystems. Location and diving data were collected concurrently, between December 2003 and February 2004, from 43 seals at Bird Island, where krill, Euphausia superba, are the main prey, and 39 at Heard Island, where mostly fish are consumed. Seals at Heard Island were shorter and lighter than those at Bird Island and they spent longer at sea, dived more frequently and spent more time in the bottom phase of dives. Generalized additive mixed effects models showed that diving behaviours differed between the islands. Both populations exploited diel vertically migrating prey species but, on average, Heard Island seals dived deeper and exceeded their estimated aerobic dive limits. We propose that the recovery of the Heard Island population may be limited by the relative inaccessibility and scarcity of food, whereas at Bird Island, the presence of abundant krill resources helps sustain extremely high numbers of seals, even with increased intra- and inter-specific competition. Both populations of fur seals appear to be constrained by their physiological limits, in terms of their optimal diving behaviour. However, there does appear to be some flexibility in strategy at the level of trip with animals adjusting their time at sea and foraging effort, in order to maximize the rate of delivery of energy to their pups.     

Quantitative analysis of aggregation behavior in juvenile blacktip sharks

Aggregation patterns of a population of juvenile blacktip sharks (Carcharhinus limbatus) within a summer nursery area were examined over a period of 3 years using an array of acoustic monitors. Individually coded acoustic tags were surgically implanted into 33–40 neonate sharks each year from 2000 to 2002 and their presence monitored continuously by the acoustic array. Data from the acoustic monitors was processed to estimate the center of activity location of each tagged shark every 30 min. Nearest neighbor analysis of shark locations revealed that sharks aggregated within the nursery in all years of the study. Sharks were aggregated most commonly during the early and late summer months (June, October and November) and became less common through the middle of the study period each year (July–September). Temporal periodicity within the data revealed a strong diel pattern with sharks aggregating during the day and dispersing at night, except in June when aggregations often persisted through the night. Aggregations were generally restricted to the mid and northern sections of the study site. Reasons for aggregations may include predator avoidance or improved feeding efficiency.     

Long‐Range Movement of Humpback Whales and Their Overlap with Anthropogenic Activity in the South Atlantic Ocean

Humpback whales (Megaptera novaeangliae) are managed by the International Whaling Commission as 7 primary populations that breed in the tropics and migrate to 6 feeding areas around the Antarctic. There is little information on individual movements within breeding areas or migratory connections to feeding grounds. We sought to better understand humpback whale habitat use and movements at breeding areas off West Africa, and during the annual migration to Antarctic feeding areas. We also assessed potential overlap between whale habitat and anthropogenic activities. We used Argos satellite‐monitored radio tags to collect data on 13 animals off Gabon, a primary humpback whale breeding area. We quantified habitat use for 3 cohorts of whales and used a state‐space model to determine transitions in the movement behavior of individuals. We developed a spatial metric of overlap between whale habitat and models of cumulative human activities, including oil platforms, toxicants, and shipping. We detected strong heterogeneity in movement behavior over time that is consistent with previous genetic evidence of multiple populations in the region. Breeding areas for humpback whales in the eastern Atlantic were extensive and extended north of Gabon late in the breeding season. We also observed, for the first time, direct migration between West Africa and sub‐Antarctic feeding areas. Potential overlap of whale habitat with human activities was the highest in exclusive economic zones close to shore, particularly in areas used by both individual whales and the hydrocarbon industry. Whales potentially overlapped with different activities during each stage of their migration, which makes it difficult to implement mitigation measures over their entire range. Our results and existing population‐level data may inform delimitation of populations and actions to mitigate potential threats to whales as part of local, regional, and international management of highly migratory marine species. Cuantificación de los Movimientos de Gran Amplitud y el Traslape Potencial con Actividad Antropogénica y las Ballenas Jorobadas en el Océano Atlántico Sur     

Movement patterns of young Caribbean reef sharks, Carcharhinus perezi, at Fernando de Noronha Archipelago, Brazil: the potential of marine protected areas for conservation of a nursery ground

The movement patterns and long-term site-fidelity of primarily juvenile Caribbean reef sharks, Carcharhinus perezi, were investigated using tag-recapture and automated telemetry at an insular nursery area, the Fernando de Noronha Archipelago, Brazil. Of the 143 externally tagged juvenile sharks (<110 cm), 22 (15.3%) were recaptured between 0 and 5 km from the site of tagging after 5–800 days at liberty, suggesting some site-fidelity in young individuals of this species. Site-fidelity and movement patterns of ten juvenile sharks ranging from 78 to 110 cm total length (TL) and one opportunistically captured adult female (224 cm TL) were also investigated for periods of up to 2 years with an array of automated telemetry receivers. Tagging and telemetry data from both inside and outside a marine protected area (MPA) show that shark abundance and activity is greatest along the part of the archipelago’s coastline least disturbed by human activity. Telemetry tracking also showed that juvenile reef sharks demonstrated a high degree of site-fidelity and occupied specific locations along the coast throughout the year, with some evidence of an increase in activity space with ontogeny. Sharks appeared to range more widely at night and there were no seasonal variations in habitat use. Our results suggest that MPAs may be a useful conservation tool to protect young C. perezi and potentially other reef-dwelling carcharhinid sharks during their early life history.     

Copepod photobehavior in a simulated natural light environment and its relation to nocturnal vertical migration

To investigate the roles of light in initiating, controlling and directing nocturnal vertical migration, photoresponses of the adult, female copepod Acartia tonsa Dana were measured under simulated natural underwater light conditions using a video system. Copepods were adapted to a range of background light levels and tested with the following stimuli: absolute quantal intensity, absolute change in quantal intensity and relative (%) change in quantal intensity. The stimulus initiating vertical movements was relative change in quantal intensity, while responsiveness was controlled by the level of light adaptation. A. tonsa swam upward in response to decreases. Response with minimal stimulation occurred at an adaptation intensity close to that in the copepod's natural habitat at the time of the migratory ascent (near the bottom of the Newport River estuary, North Carolina, near sunset). Analysis of the angles of upward movement showed that light is not a directional cue. Relative increases in intensity resulted in sinking, with minimal stimulation required at an adaptation intensity close to that in the field when the migratory descent occurs near sunrise. These results offer a reasonable explanation of how light cues may shape nocturnal vertical migratory patterns.     

Robust, comparable population metrics through collaborative photo-monitoring of whale sharks Rhincodon typus.

The formulation of conservation policy for species that are rare and migratory requires broad cooperation to ensure that adequate levels of standardized data collection are achieved and that the results of local analyses are comparable. Estimates of apparent survival rate, relative change in abundance, and proportions of newly marked and returning individuals can inform local management decisions while highlighting corresponding changes at other linked research stations. We have applied computer-assisted photo-identification and mark-recapture population modeling to whale sharks Rhincodon typus at Ningaloo Marine Park (NMP), Western Australia, to create a baseline trend for comparison with other regional aggregations of the species. We estimate several ecological parameters of interest, including an average apparent survival rate of 0.55 yr(-1) for sharks newly marked (new) and 0.83 yr(-1) for sharks captured in multiple seasons (philopatric). The average proportion of philopatric sharks is found to be 0.65 of the total population, and we derive an average population growth rate of 1.12 yr(-1) for them. Our analysis uncovered significant heterogeneity in capture and survival probabilities in this study population; our chosen model structures and data analysis account for these influences and demonstrate a good overall fit to the time-series data. The results show good correspondence between capture probability and an available measure of recapture effort, suggesting that unmodeled systematic effects contribute insignificantly to the model fits. We find no evidence of a decline in the whale shark population at NMP, and our results provide metrics of value to their future management. Overall, our study suggests an effective approach to analyzing and modeling mark-recapture data for a rare species using computer-assisted photo-identification and opportunistic data collection from ecotourism to ensure the quality and volume of data required for population analysis.     

Water column use and forage strategies of female southern elephant seals from Marion Island

The at-sea behaviour of marine top predators provides valuable insights into the distribution of prey species and strategies used by predators to exploit patchily distributed resources. We describe the water column usage and dive strategies of female southern elephant seals from Marion Island tracked between 2004 and 2008. Dives representing increases in forage effort were identified using a method that combines dive type analyses and the calculation of relative amounts of time that animals spend in the bottom phases of dives. Results from this analysis indicate that female elephant seals from Marion Island tend to display lower levels of forage effort closer to the island and display intensive opportunistic forage bouts that occur at a minimum distance of approximately 215 km from the island. Females from Marion Island dived deeper and for longer periods of time, compared to females from other populations. Most animals displayed positive diel vertical migration, evidently foraging pelagically on vertically migrating prey. A few animals displayed periods of reverse (negative) diel vertical migration, however, diving to deeper depths at night, compared to daytime. This behaviour is difficult to explain and prey species targeted during such periods unknown. Our results illustrate plasticity in foraging behaviour of southern elephant seals, as well as inter-population differences in forage strategies.     

Linking movement, diving, and habitat to foraging success in a large marine predator

Establishing where and when predators forage is essential to understanding trophic interactions, yet foraging behavior remains poorly understood in large marine carnivores. We investigated the factors leading to foraging success in gray seals (Halichoerus grypus) in the Northwest Atlantic in the first study to use simultaneous deployments of satellite transmitters, time depth recorders, and stomach-temperature loggers on a free-ranging marine mammal. Thirty-two seals were each fitted with the three types of instrumentation; however, complete records from all three instruments were obtained from only 13 individuals, underscoring the difficulty of such a multi-instrument approach. Our goal was to determine the characteristics of diving, habitat, and movement that predict feeding. We linked diving behavior to foraging success at two temporal scales: trips (days) and bouts (hours) to test models of optimal diving, which indicate that feeding can be predicted by time spent at the bottom of a dive. Using an information-theoretic approach, a Generalized Linear Mixed Model with trip duration and accumulated bottom time per day best explained the number of feeding events per trip, whereas the best predictor of the number of feeding events per bout was accumulated bottom time. We then tested whether characteristics of movement were predictive of feeding. Significant predictors of the number of feeding events per trip were angular variance (i.e., path tortuosity) and distance traveled per day. Finally, we integrated measures of diving, movement, and habitat at four temporal scales to determine overall predictors of feeding. At the 3-h scale, mean bottom time and distance traveled were the most important predictors of feeding frequency, whereas at the 6-h and 24-h time scales, distance traveled alone was most important. Bathymetry was the most significant predictor of feeding at the 12-h interval, with feeding more likely to occur at deeper depths. Our findings indicate that several factors predict feeding in gray seals, but predictor variables differ across temporal scales such that environmental variation becomes important at some scales and not others. Overall, our results illustrate the value of simultaneously recording and integrating multiple types of information to better understand the circumstances leading to foraging success.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

Mechanisms to initiate a Heterosigma akashiwo red tide in Osaka bay

The ecological role of diel vertical migration of Heterosigma akashiwo Hada to initiate red tide was investigated in Tanigawa Fishing Port and Sano Harbor, Osaka Bay, Japan during red tide seasons in 1979 and 1980. This species migrated toward the surface early in the morning at a velocity of 1.0 to 1.3 m h^-1. Downward migration was found in the afternoon, and more than 2.0×10³ cells ml^-1 aggregated in the bottom layer at night. The upward migration started before sunrise and downward shifting occurred prior to sunset. The movement is presumably associated with circadian rhythm, which is known as one of the biological periodicities. H. akashiwo crossed steep temperature and salinity gradients (6.5°C and 5.7 S) during the diel vertical migration. High values of particulate organic carbon and nitrogen concentrations were obtained in dialysis bags suspended in situ at identical layers with high cell concentration, while the values for surface and bottom bags were comparatively low. The results reveal that H. akashiwo migrates toward the sea surface to carry out photosynthesis effectively, and to the bottom to utilize nutrients efficiently.     

Vertical migration strategies with respect to advection and stratification in a semi-enclosed lough: a comparison of mero- and holozooplankton

Patterns of zooplankton vertical movement are often difficult to interpret because of multiple, complex and confounding environmental factors. Behavioural adaptations to these environmental variables are compared within and between the holo- and meroplankton constituents of a community. We used a nested design to analyse patterns at several scales in time; (semi-diel, diel, spring-neap tidal cycle and season) and two in space; (depth and site). To reduce complexity and aid interpretation we studied a semi-isolated community in a semi-enclosed, seasonally stratified sea lough (Lough Hyne Marine Nature Reserve, Ireland). In this, the main environmental gradient was water flow rate (or water residence time) caused by tidal currents. Vertical profiles of abundance showed that populations of the most abundant species of holo- and meroplankton in the lough have considerable behavioural plasticity, enabling them to switch between sedentary and migratory behaviour and patterns of migration. Some species migrate vertically in synchrony with diel cycles and others in response to semi-diel tidal currents; a few do both, but the majority did neither. It is suggested that water column structure and hydrographic discontinuities caused by flow rate and pycnocline dynamics are responsible for the variable patterns of vertical migration and distribution.Communicated by J.P. Thorpe, Port Erin     

Satellite tagging of blue sharks (Prionace glauca) and other pelagic sharks off eastern Australia: depth behaviour, temperature experience and movements

Satellite telemetry was used to study the movements and behaviour of ten blue sharks and one individual each of shortfin mako, thresher and bigeye thresher off eastern Australia. The tracks showed latitudinal movements of up to 1,900 km, but none of the sharks travelled away from the eastern Australian region. Tracking periods did not exceed 177 days. All species showed oscillatory dive behaviour between the surface layers to as deep as 560–1,000 m. Blue sharks spent 35–58% of their time in <50 m depths and 10–16% of their time in >300 m. Of these four species, the bigeye thresher spent the least time in the surface layers and the most time at >300 m depth. All four species showed clear diel behaviour generally occupying shallower depths at night than during the day. Blue sharks were mainly in 17.5–20.0°C water, while the thresher sharks showed a more bimodal temperature distribution.     

Movement and environmental preferences of Greenland sharks (<Emphasis Type="Italic">Somniosus microcephalus</Emphasis>) electronically tagged in the St. Lawrence Estuary,Canada

Three Greenland sharks (Somniosus microcephalus) were tagged with electronic tags, in Baie St. Pancrace, St. Lawrence Estuary, Quebec, Canada. One shark was tagged on 23 July 2004, with an acoustic telemetry tag. Two sharks were each tagged with a pop-up satellite archival tag (PSAT) on 27 August 2004. Two of the sharks remained in or close to the bay, one for 47 days and the other for at least 66 days. The third shark left the bay immediately after tagging on 27 August 2004. This shark entered the main channel of the St. Lawrence Estuary, and had moved 114.9 km upstream by 1 November 2004 when the tag reported to ARGOS satellites. The tags provided a total of 179 days of data on the movement and environmental preferences of Greenland sharks in the St. Lawrence Estuary. Sharks that reported depth and ambient water temperature data from the bay showed significant diel differences in depth preferences and corresponding ambient temperatures. The sharks remained near the bottom of the water column during the day and displayed increased vertical movements at night. The shark that resided in the main channel did not show this pattern, but generally remained at depths between 325 and 352 m. Sharks in the bay experienced water temperatures that ranged from −1.1 to 8.6°C at depths from 0 to 67 m. In the main channel the shark experienced temperatures that ranged from 1.0 to 5.4°C at depths from 132 to 352 m. This is the first report of numerous Greenland sharks inhabiting shallow near shore bays during summer and autumn.