Sascha K. Hooker

Marine Reserves as a Tool for Ecosystem-Based Management: The Potential Importance of Megafauna

Abstract Marine predators attract significant attention in ocean conservation planning and are therefore often used politically to promote reserve designation. We discuss whether their ecology and life history can help provide a rigorous ecological foundation for marine reserve design. In general, we find that reserves can benefit marine megafauna, and that megafauna can help establish target areas and boundaries for ecosystem reserves. However, the spatial nature of the interplay between potential threats and predator life histories requires careful consideration for the establishment of effective reserves. Modeling tools such as demographic sensitivity analysis will aid in establishing protection for different life stages and distributional ranges. The need for pelagic marine reserves is becoming increasingly apparent, and it is in this venue that marine predators may be most effectively used as indicator species of underlying prey distribution and ecosystem processes.

Deadly diving? Physiological and behavioural management of decompression stress in diving mammals

Decompression sickness (DCS; ‘the bends’) is a disease associated with gas uptake at pressure. The basic pathology and cause are relatively well known to human divers. Breath-hold diving marine mammals were thought to be relatively immune to DCS owing to multiple anatomical, physiological and behavioural adaptations that reduce nitrogen gas (N2) loading during dives. However, recent observations have shown that gas bubbles may form and tissue injury may occur in marine mammals under certain circumstances. Gas kinetic models based on measured time-depth profiles further suggest the potential occurrence of high blood and tissue N2 tensions. We review evidence for gas-bubble incidence in marine mammal tissues and discuss the theory behind gas loading and bubble formation. We suggest that diving mammals vary their physiological responses according to multiple stressors, and that the perspective on marine mammal diving physiology should change from simply minimizing N2 loading to management of the N2 load. This suggests several avenues for further study, ranging from the effects of gas bubbles at molecular, cellular and organ function levels, to comparative studies relating the presence/absence of gas bubbles to diving behaviour. Technological advances in imaging and remote instrumentation are likely to advance this field in coming years.

Estimating the effect of lung collapse and pulmonary shunt on gas exchange during breath-hold diving: The Scholander and Kooyman legacy

We developed a mathematical model to investigate the effect of lung compression and collapse (pulmonary shunt) on the uptake and removal of O(2), CO(2) and N(2) in blood and tissue of breath-hold diving mammals. We investigated the consequences of pressure (diving depth) and respiratory volume on pulmonary shunt and gas exchange as pressure compressed the alveoli. The model showed good agreement with previous studies of measured arterial O(2) tensions (Pa(O)(2)) from freely diving Weddell seals and measured arterial and venous N(2) tensions from captive elephant seals compressed in a hyperbaric chamber. Pulmonary compression resulted in a rapid spike in Pa(O)(2) and arterial CO(2) tension, followed by cyclical variation with a periodicity determined by Q(tot). The model showed that changes in diving lung volume are an efficient behavioural means to adjust the extent of gas exchange with depth. Differing models of lung compression and collapse depth caused major differences in blood and tissue N(2) estimates. Our integrated modelling approach contradicted predictions from simple models, and emphasised the complex nature of physiological interactions between circulation, lung compression and gas exchange. Overall, our work suggests the need for caution in interpretation of previous model results based on assumed collapse depths and all-or-nothing lung collapse models.

Salinity sensors on seals: use of marine predators to carry CTD data loggers

Diving marine predators have been used to collect data on ocean temperature, but salinity measurements have not previously been incorporated into predator-borne data loggers. Here we present data on initial calibration and field trials of a new conductivity, temperature and depth (CTD) data logger used alongside a satellite-positioning transmitter to provide three-dimensional oceanographic information. This provides CTD data analogous to that collected by a ship-deployed undulating oceanographic recorder. Calibration tests of these units showed a near-field effect caused by the proximity of material to the tag, but demonstrate that the resulting data offset can be removed by post hoc calibration. Field tests of the system were conducted on 16 female Antarctic fur seals (Arctocephalus gazella) at Bird Island, South Georgia. These results matched those found by standard ship-based survey techniques, but suggest temporal variability in the structure and location of the two water masses found to the north of South Georgia. Overall, this initial proof-of-concept work is encouraging; future refinement of this technique is likely to provide an additional data source for both oceanographers and biologists.

Could beaked whales get the bends? Effect of diving behaviour and physiology on modelled gas exchange for three species: Ziphius cavirostris, Mesoplodon densirostris and Hyperoodon ampullatus.

A mathematical model, based on current knowledge of gas exchange and physiology of marine mammals, was used to predict blood and tissue tension N2 (P(N2)) using field data from three beaked whale species: northern bottlenose whales, Cuviers beaked whales, and Blainvilles beaked whales. The objective was to determine if physiology (body mass, diving lung volume, dive response) or dive behaviour (dive depth and duration, changes in ascent rate, diel behaviour) would lead to differences in P(N2) levels and thereby decompression sickness (DCS) risk between species. Diving lung volume and extent of the dive response had a large effect on end-dive P(N2). The dive profile had a larger influence on end-dive P(N2) than body mass differences between species. Despite diel changes in dive behaviour, P(N2) levels showed no consistent trend. Model output suggested that all three species live with tissue P(N2) levels that would cause a significant proportion of DCS cases in terrestrial mammals. Cuviers beaked whale diving behaviour appears to put them at higher risk than the other species, which may explain their prevalence in strandings after the use of mid-frequency sonar.

Using Energetic Models to Investigate the Survival and Reproduction of Beaked Whales (family Ziphiidae)

Mass stranding of several species of beaked whales (family Ziphiidae) associated with exposure to anthropogenic sounds has raised concern for the conservation of these species. However, little is known about the species’ life histories, prey or habitat requirements. Without this knowledge, it becomes difficult to assess the effects of anthropogenic sound, since there is no way to determine whether the disturbance is impacting the species’ physical or environmental requirements. Here we take a bioenergetics approach to address this gap in our knowledge, as the elusive, deep-diving nature of beaked whales has made it hard to study these effects directly. We develop a model for Ziphiidae linking feeding energetics to the species’ requirements for survival and reproduction, since these life history traits would be the most likely to be impacted by non-lethal disturbances. Our models suggest that beaked whale reproduction requires energy dense prey, and that poor resource availability would lead to an extension of the inter-calving interval. Further, given current information, it seems that some beaked whale species require relatively high quality habitat in order to meet their requirements for survival and reproduction. As a result, even a small non-lethal disturbance that results in displacement of whales from preferred habitats could potentially impact a population if a significant proportion of that population was affected. We explored the impact of varying ecological parameters and model assumptions on survival and reproduction, and find that calf and fetus survival appear more readily affected than the survival of adult females.

Ascent exhalations of Antarctic fur seals: a behavioural adaptation for breath-hold diving?

Novel observations collected from video, acoustic and conductivity sensors showed that Antarctic fur seals consistently exhale during the last 50–85% of ascent from all dives (10–160 m, n > 8000 dives from 50 seals). The depth of initial bubble emission was best predicted by maximum dive depth, suggesting an underlying physical mechanism. Bubble sound intensity recorded from one seal followed predictions of a simple model based on venting expanding lung air with decreasing pressure. Comparison of air release between dives, together with lack of variation in intensity of thrusting movement during initial descent regardless of ultimate dive depth, suggested that inhaled diving lung volume was constant for all dives. The thrusting intensity in the final phase of ascent was greater for dives in which ascent exhalation began at a greater depth, suggesting an energetic cost to this behaviour, probably as a result of loss of buoyancy from reduced lung volume. These results suggest that fur seals descend with full lung air stores, and thus face the physiological consequences of pressure at depth. We suggest that these regular and predictable ascent exhalations could function to reduce the potential for a precipitous drop in blood oxygen that would result in shallow–water blackout.

First indications that northern bottlenose whales are sensitive to behavioural disturbance from anthropogenic noise

Although northern bottlenose whales were the most heavily hunted beaked whale, we have little information about this species in its remote habitat of the North Atlantic Ocean. Underwater anthropogenic noise and disruption of their natural habitat may be major threats, given the sensitivity of other beaked whales to such noise disturbance. We attached dataloggers to 13 northern bottlenose whales and compared their natural sounds and movements to those of one individual exposed to escalating levels of 1–2 kHz upsweep naval sonar signals. At a received sound pressure level (SPL) of 98 dB re 1 μPa, the whale turned to approach the sound source, but at a received SPL of 107 dB re 1 μPa, the whale began moving in an unusually straight course and then made a near 180° turn away from the source, and performed the longest and deepest dive (94 min, 2339 m) recorded for this species. Animal movement parameters differed significantly from baseline for more than 7 h until the tag fell off 33–36 km away. No clicks were emitted during the response period, indicating cessation of normal echolocation-based foraging. A sharp decline in both acoustic and visual detections of conspecifics after exposure suggests other whales in the area responded similarly. Though more data are needed, our results indicate high sensitivity of this species to acoustic disturbance, with consequent risk from marine industrialization and naval activity.

Data sampling options for animal-borne video cameras: considerations based on deployments with antarctic fur seals

Although the use of animal-borne video cameras has brought great insight into the lives of animals, there are still several limitations in using them to record observational data. The ethical demand for reduced camera size limits battery capacity, which thus restricts recording duration. In addition to design considerations, there are several sampling options available for extending sampling duration following tag deployment. These options include the use of duty cycling (for sampling at preset times), a depth trigger and varying the sampling interval between images. We investigate the consequences of these options based on foraging data collected from Antarctic fur seals using the Venus camera system (Wild Insight Ltd). We show the effect of differing sampling protocols on the resulting data. By sub-sampling our dataset, we show how changes in sampling protocol affect the number of prey images collected and the measurement of simple foraging variables such as the time spent feeding. In addition, we ran simulations based on altering the foraging parameters: prey-encounter rate (waiting time) and handling time (feeding event duration), to more broadly investigate how changes in sampling frequency would impact data collection for species with varying foraging characteristics. We show that, irrespective of sampling frequency, the proportion of prey images recorded remains constant, suggesting that coverage of a foraging trip could be extended for investigation of time-invariant processes by using an increased sampling interval. Measurement of rates or durations of behavioral events (such as prey-encounter rate or handling time), however, requires higher sampling rates to increase the precision of parameter values.

Pilot cetacean survey of the sub-Arctic North Atlantic utilizing a cruise-ship platform

Using a large passenger cruise-ship (MV ‘Discovery’) as a platform, a cetacean survey was conducted between 1 August and 3 September 2005 across the northern North Atlantic Ocean and back, covering waters between the UK, Iceland, Greenland and Canada. The objective was to collect sightings data for all cetacean species encountered to begin to collate information for a large-scale sightings database. Two observers employed standard-distance sampling techniques and visual observations (naked eye) to scan for cetaceans during daylight hours and favourable weather conditions. Approximately 112 h were spent surveying over 23 d. A total of 13 cetacean species were encountered, with 173 separate sightings recorded, totalling over 2000 animals. The most commonly sighted species were minke whales (Balaenoptera acutorostrata: N=44), Atlantic white-sided dolphins (Lagenorhynchus acutus: N=34, with group sizes of up to 600), and humpback whales (Megaptera novaeangliae: N=31). The distribution of minke whales, Atlantic white-sided dolphins and fin whales was linked to underlying oceanographic variables. Overall patterns of distribution were consistent with our understanding of the summer distributions for the species encountered. The survey highlighted the cetacean diversity of the northern North Atlantic region. Repeat work using this methodology will allow an examination of species’ distributions and estimation of their relative abundance.