Jayson M. Semmens

Automated acoustic tracking of aquatic animals: scales, design and deployment of listening station arrays

The recent introduction of low-cost, moored data-logging acoustic receivers has provided opportunities for tracking marine organisms over small (hundreds of metres) and large scales (hundreds of kilometres). Acoustic receivers have been deployed in many different environments to examine specific hypotheses regarding the movement of aquatic species. This technology provides many advantages for studying aquatic animal movement patterns, but also has limitations and provides unique difficulties for users. Study design, applications, advantages and limitations are discussed with examples from past and current studies. Data management and analysis techniques are in their infancy and few standardised techniques exist. Complications with data management and potential data analysis techniques are discussed. Examples from the literature are utilised wherever possible to provide useful references.

Understanding octopus growth: patterns, variability and physiology

Octopuses are generally characterised by rapid non-asymptotic growth, with high individual variability. However, in situ octopus growth is not well understood. The lack of an ageing method has resulted in the majority of our understanding of octopus growth coming from laboratory studies. Despite not being applicable to cephalopods, Modal Progression Analysis (MPA) of length-frequency data is the most common method for examining in situ octopus growth. Recently, counting growth increments in beaks and vestigial shells, and quantifying lipofuscin in brain tissue, have all shown promise for the ageing octopus. Octopuses generally demonstrate two-phase growth in the laboratory, with physiological changes possibly associated with the switch between an initial rapid exponential phase and a slower power growth phase. Temperature and food ration and quality are key factors influencing the initial growth phase. Temperature, however, does not appear to affect the second phase in any consistent way, perhaps because maturity stage can influence the growth response. There may be basic differences in the mechanisms of octopus muscle growth compared with that of other cephalopods. Furthermore, higher relative maintenance energy expenditure, along with the low energy content of their prey, may account for the relatively slow growth of deep-sea octopuses compared to littoral species.

Approaches to resolving cephalopod movement and migration patterns

Cephalopod movement occurs during all phases of the life history, with the abundance and location of cephalopod populations strongly influenced by the prevalence and scale of their movements. Environmental parameters, such as sea temperature and oceanographic processes, have a large influence on movement at the various life cycle stages, particularly those of oceanic squid. Tag recapture studies are the most common way of directly examining cephalopod movement, particularly in species which are heavily fished. Electronic tags, however, are being more commonly used to track cephalopods, providing detailed small- and large-scale movement information. Chemical tagging of paralarvae through maternal transfer may prove to be a viable technique for tracking this little understood cephalopod life stage, as large numbers of individuals could be tagged at once. Numerous indirect methods can also be used to examine cephalopod movement, such as chemical analyses of the elemental and/or isotopic signatures of cephalopod hard parts, with growing interest in utilising these techniques for elucidating migration pathways, as is commonly done for fish. Geographic differences in parasite fauna have also been used to indirectly provide movement information, however, explicit movement studies require detailed information on parasite-host specificity and parasite geographic distribution, which is yet to be determined for cephalopods. Molecular genetics offers a powerful approach to estimating realised effective migration rates among populations, and continuing developments in markers and analytical techniques hold the promise of more detailed identification of migrants. To date genetic studies indicate that migration in squids is extensive but can be blocked by major oceanographic features, and in cuttlefish and octopus migration is more locally restricted than predictions from life history parameters would suggest. Satellite data showing the location of fishing lights have been increasingly used to examine the movement of squid fishing vessels, as a proxy for monitoring the movement of the squid populations themselves, allowing for the remote monitoring of oceanic species.

In situ measurement of coastal ocean movements and survival of juvenile Pacific salmon.

Many salmon populations in both the Pacific and Atlantic Oceans have experienced sharply decreasing returns and high ocean mortality in the past two decades, with some populations facing extirpation if current marine survival trends continue. Our inability to monitor the movements of marine fish or to directly measure their survival precludes experimental tests of theories concerning the factors regulating fish populations, and thus limits scientific advance in many aspects of fisheries management and conservation. Here we report a large-scale synthesis of survival and movement rates of free-ranging juvenile salmon across four species, 13 river watersheds, and 44 release groups of salmon smolts (>3,500 fish tagged in total) in rivers and coastal ocean waters, including an assessment of where mortality predominantly occurs during the juvenile migration. Of particular importance, our data indicate that, over the size range of smolts tagged, (i) smolt survival was not strongly related to size at release, (ii) tag burden did not appear to strongly reduce the survival of smaller animals, and (iii) for at least some populations, substantial mortality occurred much later in the migration and more distant from the river of origin than generally expected. Our findings thus have implications for determining where effort should be invested to improve the accuracy of salmon forecasting, to understand the mechanisms driving salmon declines, and to predict the impact of climate change on salmon stocks.

Accelerometry estimates field metabolic rate in giant Australian cuttlefish Sepia apama during breeding

1. Estimating the metabolic rate of animals in nature is central to understanding the physiological, behavioural and evolutionary ecology of animals. Doubly labelled water and heart-rate methods are the most commonly used approaches, but both have limitations that preclude their application to some systems. 2. Accelerometry has emerged as a powerful tool for estimating energy expenditure in a range of animals, but is yet to be used to estimate field metabolic rate in aquatic taxa. We combined two-dimensional accelerometry and swim-tunnel respirometry to estimate patterns of energy expenditure in giant Australian cuttlefish Sepia apama during breeding. 3. Both oxygen consumption rate (Vo2) and swimming speed showed strong positive associations with body acceleration, with coefficients of determination comparable to those using similar accelerometers on terrestrial vertebrates. Despite increased activity during the day, field metabolic rate rarely approached Vo2, and night-time Vo2 was similar to that at rest. 4. These results are consistent with the life-history strategy of this species, which has a poor capacity to exercise anaerobically, and a mating strategy that is visually based. With the logistical difficulties associated with observation in aquatic environments, accelerometry is likely to prove a valuable tool for estimating energy expenditure in aquatic animals.

Site fidelity and sex-specific migration in a mobile apex predator: implications for conservation and ecosystem dynamics

Combining movement behaviour with other ecological information of predators and their prey is essential for an adequate understanding of ecosystem dynamics. The movement patterns of broadnose sevengill sharks, Notorynchus cepedianus, were monitored with acoustic and satellite technology in coastal areas of southeast Tasmania, Australia. Individuals were tagged in two habitats (Norfolk Bay and the Derwent Estuary) for which we had ecological information such as diet, population structure and abundance. Notorynchus cepedianus showed seasonal site fidelity in the use of the coastal habitats. The general pattern was for sharks to exit coastal areas over winter and females to return the following spring and males in summer. Their movement into these coastal areas coincided with high seasonal abundance of their known prey species during summer, suggesting feeding site fidelity. Individuals tagged in two coastal areas showed low spatial and dietary overlap, suggesting localized site fidelity and fine spatial scale resource partitioning. This has rarely been reported for large mobile predators. Both satellite and acoustic methods showed that males make northerly migrations during winter to distances of at least 1000 km. The combined use of tracking, diet and abundance information demonstrated that N. cepedianus are likely to exert significant predation pressure on prey inhabiting these areas during summer. Overall, this study highlights the benefit of complementing movement data with other ecological information to understand the habitat use of large mobile predators and their potential influences on ecosystem structure and function.

AN EXAMINATION OF THE ROLE OF THE DIGESTIVE GLAND OF TWO LOLIGINID SQUIDS,WITH RESPECT TO LIPID : STORAGE OR EXCRETION ?

The role of the digestive gland, with respect to non–structural lipid, was examined using proximal analysis, histochemistry and quantitative histological techniques in the tropical loliginid squids Sepioteuthis lessoniana (Lesson) and Photololigo sp. The digestive gland of both species was characterized by large and numerous lipid droplets in the apical portion of the digestive cells and very few in the basal portion. The apical lipid droplets were released into the lumen of the gland and subsequently rapidly removed. Despite the numerous large apical lipid droplets, the lipid concentration in the digestive glands of S. lessoniana and Photololigo sp. was lower than that reported for most squid species. There was no relationship between lipid concentration and stage of digestion, suggesting that lipid is not stored in the gland after a meal. There was also no relationship between lipid concentration and the sex of an individual or stage of reproductive maturity, suggesting that these squids are not storing lipid in the digestive gland for use in fuelling reproductive maturation or providing an energy source for oocytes. I believe this study is the first to combine proximal analysis and quantitative histological techniques to examine the role of the squid digestive gland with respect to non–structural lipids. The results indicate that the digestive gland of these tropical loliginid squids is excreting, not storing, excess dietary lipid.

Rain reverses diel activity rhythms in an estuarine teleost

Activity rhythms are ubiquitous in nature, and generally synchronized with the day–night cycle. Several taxa have been shown to switch between nocturnal and diurnal activity in response to environmental variability, and these relatively uncommon switches provide a basis for greater understanding of the mechanisms and adaptive significance of circadian (approx. 24 h) rhythms. Plasticity of activity rhythms has been identified in association with a variety of factors, from changes in predation pressure to an altered nutritional or social status. Here, we report a switch in activity rhythm that is associated with rainfall. Outside periods of rain, the estuarine-associated teleost Acanthopagrus australis was most active and in shallower depths during the day, but this activity and depth pattern was reversed in the days following rain, with diurnality restored as estuarine conductivity and turbidity levels returned to pre-rain levels. Although representing the first example of a rain-induced reversal of activity rhythm in an aquatic animal of which we are aware, our results are consistent with established models on the trade-offs between predation risk and foraging efficiency.

Fine-Scale Movements of the Broadnose Sevengill Shark and Its Main Prey, the Gummy Shark

Information on the fine-scale movement of predators and their prey is important to interpret foraging behaviours and activity patterns. An understanding of these behaviours will help determine predator-prey relationships and their effects on community dynamics. For instance understanding a predators movement behaviour may alter pre determined expectations of prey behaviour, as almost any aspect of the preys decisions from foraging to mating can be influenced by the risk of predation. Acoustic telemetry was used to study the fine-scale movement patterns of the Broadnose Sevengill shark Notorynchus cepedianus and its main prey, the Gummy shark Mustelus antarcticus, in a coastal bay of southeast Tasmania. Notorynchus cepedianus displayed distinct diel differences in activity patterns. During the day they stayed close to the substrate (sea floor) and were frequently inactive. At night, however, their swimming behaviour continually oscillated through the water column from the substrate to near surface. In contrast, M. antarcticus remained close to the substrate for the entire diel cycle, and showed similar movement patterns for day and night. For both species, the possibility that movement is related to foraging behaviour is discussed. For M. antarcticus, movement may possibly be linked to a diet of predominantly slow benthic prey. On several occasions, N. cepedianus carried out a sequence of burst speed events (increased rates of movement) that could be related to chasing prey. All burst speed events during the day were across the substrate, while at night these occurred in the water column. Overall, diel differences in water column use, along with the presence of oscillatory behaviour and burst speed events suggest that N. cepedianus are nocturnal foragers, but may opportunistically attack prey they happen to encounter during the day.

Global proliferation of cephalopods

Human activities have substantially changed the worlds oceans in recent decades, altering marine food webs, habitats and biogeochemical processes [1]. Cephalopods (squid, cuttlefish and octopuses) have a unique set of biological traits, including rapid growth, short lifespans and strong life-history plasticity, allowing them to adapt quickly to changing environmental conditions [2-4]. There has been growing speculation that cephalopod populations are proliferating in response to a changing environment, a perception fuelled by increasing trends in cephalopod fisheries catch [4,5]. To investigate long-term trends in cephalopod abundance, we assembled global time-series of cephalopod catch rates (catch per unit of fishing or sampling effort). We show that cephalopod populations have increased over the last six decades, a result that was remarkably consistent across a highly diverse set of cephalopod taxa. Positive trends were also evident for both fisheries-dependent and fisheries-independent time-series, suggesting that trends are not solely due to factors associated with developing fisheries. Our results suggest that large-scale, directional processes, common to a range of coastal and oceanic environments, are responsible. This study presents the first evidence that cephalopod populations have increased globally, indicating that these ecologically and commercially important invertebrates may have benefited from a changing ocean environment.