Biosonar behaviour of free-ranging porpoises

Detecting objects in their paths is a fundamental perceptional function of moving organisms. Potential risks and rewards, such as prey, predators, conspecifics or non-biological obstacles, must be detected so that an animal can modify its behaviour accordingly. However, to date few studies have considered how animals in the wild focus their attention. Dolphins and porpoises are known to actively use sonar or echolocation. A newly developed miniature data logger attached to a porpoise allows for individual recording of acoustical search efforts and inspection distance based on echolocation. In this study, we analysed the biosonar behaviour of eight free-ranging finless porpoises (Neophocaena phocaenoides) and demonstrated that these animals inspect the area ahead of them before swimming silently into it. The porpoises inspected distances up to 77 in, whereas their swimming distance without using sonar was less than 20 in. The inspection distance was long enough to ensure a wide safety margin before facing real risks or rewards. Once a potential prey item was detected, porpoises adjusted their inspection distance from the remote target throughout their approach.

Soft sediment ecology of Lough Hyne Marine Reserve - spatial and temporal patterns

Distribution of soft sediment benthic fauna and the environmental factors affecting them were studied, to investigate changes across spatial and temporal scales. Investigations took place at Lough Hyne Marine Reserve using a range of methods. Data on the sedimentation rates of organic and inorganic matter were collected at monthly intervals for one year at a number of sites around the Lough, by use of vertical midwater-column sediment traps. Sedimentation of these two fractions were not coupled; inorganic matter sedimentation depended on hydrodynamic and weather factors, while the organic matter sedimentation was more complex, being dependent on biological and chemical processes in the water column. The effects of regular hypoxic episodes on benthic fauna due to a natural seasonal thermocline were studied in the deep Western Trough, using camera-equipped remotely-operated vehicle to follow transects, on a three-monthly basis over one year. In late summer, the area below the thermocline of the Western Trough was devoid of visible fauna. Decapod crustaceans were the first taxon to make use of ameliorating oxygen conditions in autumn, by darting below the thermocline depth, most likely to scavenge. This was indicated by tracks that they left on the surface of the Trough floor. Some species, most noticeably Fries’ goby Lesueurigobius friesii, migrated below the thermocline depth when conditions were normoxic and established semi-permanent burrows. Their population encompassed all size classes, indicating that this habitat was not limited to juveniles of this territorial species. Recolonisation by macrofauna and burrowing megafauna was studied during normoxic conditions, from November 2009 to May 2010. Macrofauna displayed a typical post-disturbance pattern of recolonisation with one species, the polychaete Scalibregma inflatum, occurring at high abundance levels in March 2010. In May, this population had become significantly reduced and a more diverse community was established. The abundance of burrowing infauna comprising decapods crabs and Fries’ gobies, was estimated by identifying and counting their distinctive burrow structures. While above the summer thermocline depth, burrow abundance increased in a linear fashion, below the thermocline depth a slight reduction of burrow abundance occurred in May, when oxygen conditions deteriorated again. The majority of the burrows occurring in May were made by Fries’ gobies, which are thought to encounter low oxygen concentrations in their burrows. Reduction in burrow abundance of burrowing shrimps Calocaris macandreae and Callianassa subterranea (based on descriptions of burrow structures from the literature), from March to May, might be related to their reduced activity in hypoxia, leading to loss of structural burrow maintenance. Spatial and temporal changes to macrofaunal assemblage structures were studied seasonally for one year across 5 sites in the Lough and subject to multivariate statistical analysis. Assemblage structures were significantly correlated with organic matter levels in the sediment, the amounts of organic matter settling out of the water column one month before macrofaunal sampling took place as well as current speed and temperature. This study was the first to investigate patterns and processes in the Lough soft sediment ecology across all 3 basins on a temporal and spatial scale. An investigation into the oceanographic aspects of the development, behaviour and break-down of the summer thermocline of Lough Hyne was performed in collaboration with researchers from other Irish institutions.

The life history and ecology of black scabbardfish (Aphanopus carbo Lowe 1839) in the north-east Atlantic

The black scabbardfish is a deep water species that supports commercial fisheries across a large area of the NE Atlantic shelf. The life history of black scabbardfish is poorly understood and a major unresolved issue is population structure. In this study it was used a combination of methodologies to get further knowledge in the life history and population structure of A. carbo over its wide distribution range in the Northeast Atlantic. The new knowledge acquired during this study, will increase our ability to better manage this species in the NE Atlantic. It has been postulated that fish caught to the west of the British Isles are pre-adults that migrate further south (to Madeira) for spawning, implying a single panmictic population. In this study, specimens of Aphanopus carbo were sampled between September 2008 and May 2010 from two different areas: NW Scotland (French trawlers and deep water surveys) and Madeira Islands (longliners commercial landings). Geographical differences in reproductive state of scabbardfish were evident, supportive of a north-south migration theory. In the northern area, all specimens found were immature, while in Madeira all maturity stages were observed. In Madeira, spawning occurred during the fourth quarter, with peak maturity in October (males) and in November (females). The age of this species has proven difficult and has led to different and contradictory age and growth estimates. For this study, we used two reading interpretations to determine age and estimate the growth parameters. To the west of the British Isles, specimens reached a lower maximum age and had a higher growth rate than those caught off Madeira. These differences are consistent with the theory of a single population of black scabbardfish in the NE Atlantic, highly segregate, with smaller, immature and younger fish caught to the west of the British Isles and bigger and mature caught in Madeira Islands. The feeding ecology showed strong evidence that the diet of black scabbardfish is associated with the spawning migration of blue whiting, which may support a northerly feeding migration theory for black scabbardfish. The stable isotope analyses in the muscle of black scabbardfish identified that black scabbardfish feeds on species with epipelagic and benthopelagic affinities. Comparison with stable isotope analysis in Madeira samples indicated that black scabbardfish feed at a similar trophic level and has the same trophic niche width in both areas, assuming similar baseline isotope compositions. Otolith stable isotopes (oxygen - δ18O and nitrogen - δ15N) analyses were used as a tool to clarify migratory behaviour. Otolith isotope ratios can provide insight into whether adults caught around Madeira fed in an isotopically depleted northerly ecosystem (NW Scotland) during their pre-adult period and then migrate towards south to spawn. Overall, the results support a south-north migration of pre adult fish from spawning areas around Madeira and a north-south migration from the west of Scotland to the spawning areas. Given its life cycle there is an urgent need that the management process recognizes the existence of a continuous widely distributed stock of black scabbardfish between the west of the British Isles and Madeira. The results highlight large scale dispersal in this species which needs to be treated as a highly migratory species and be managed as a single population.

Ecology of frontier populations of the invasive grey squirrel (Sciurus carolinensis) in Ireland

The rise in invasive species, together with habitat destruction, is associated with worldwide declines in biodiversity and ecosystem functioning. Management of invasive species, as well as amelioration of invasion impacts, provide challenges to species and ecosystem ecologists and conservation managers. Although any species can become invasive if it is transported to, establishes in and spreads in a new environment outside of its native range, rodents are a particularly frequent invader. Rodent introductions are often inadvertent but are also commonly intentional as these animals are traded and transported as pets and may escape from captivity. Tree squirrel species are attractive to humans and are able to establish populations with only a few founding individuals, making them a group well suited to performing the role of biological invaders. The eastern grey squirrel (Sciurus carolinensis) is the most commonly introduced squirrel species worldwide. This research addressed the grey squirrel invasion and frontier population biology. Novel results were generated through diverse research techniques. Public sighting surveys and hairtube surveys were used to locate the southern frontier of grey squirrel range expansion in Ireland. A 22-month intensive live trapping study of two frontier populations facilitated the collection of personality and demographic data from squirrels in increasing populations. A systematic literature search on grey squirrel demography provided context for the studied populations, among frontier and established introduced populations, as well as those in the native range. Advanced spatially explicit population modeling techniques predicted future range expansion and objectively compared the outcomes of 12 grey squirrel management strategies. The methods and results are discussed in both a basic scientific and applied invasion management context. An improved understanding of the behaviour, population dynamics, and future scenarios at the frontier of species invasions is crucial for managers worldwide and this is provided here for the grey squirrel in Ireland.

Freezing behaviour facilitates bioelectric crypsis in cuttlefish faced with predation risk.

Cephalopods, and in particular the cuttlefish Sepia officinalis, are common models for studies of camouflage and predator avoidance behaviour. Preventing detection by predators is especially important to this group of animals, most of which are soft-bodied, lack physical defences, and are subject to both visually and non-visually mediated detection. Here, we report a novel cryptic mechanism in S. officinalis in which bioelectric cues are reduced via a behavioural freeze response to a predator stimulus. The reduction of bioelectric fields created by the freeze-simulating stimulus resulted in a possible decrease in shark predation risk by reducing detectability. The freeze response may also facilitate other non-visual cryptic mechanisms to lower predation risk from a wide range of predator types.

Flexible diel vertical migration behaviour of zooplankton in the Irish Sea

The diel vertical migration (DVM) of the whole plankton community was investigated in the central and coastal Irish Sea. Generally, more than 60% of the plankton did not perform significant DVM. A correlation analysis of the weighted mean depths of different organisms and their potential predators suggested relationships between two groups, Oithona spp., copepod nauplii and fish larvae, and between Pseudocalanus elongatus, Calanus spp. and chaetognaths. The organisms showing significant DVM were chaetognaths (Sagitta spp.), Calanus spp. and P. elongatus. Calanus spp. showed clear ontogenic variations in DVM, and along with P. elongatus demonstrated great flexibility both in the amplitude and direction of migration. P. elongatus did not migrate in the coastal area and Calanus spp. showed a clear reverse migration. The direction of migration appeared to be related to the vertical position of the chaetognaths in the water column during the day.

Seasonal movements and behaviour of basking sharks from archival tagging: No evidence of winter hibernation

Habitat selection processes in highly migratory animals such as sharks and whales are important to understand because they influence patterns of distribution, availability and therefore catch rates. However, spatial strategies remain poorly understood over seasonal scales in most species, including, most notably, the plankton-feeding basking shark Cetorhinus maximus. It was proposed nearly 50 yr ago that this globally distributed species migrates from coastal summer-feeding areas of the northeast Atlantic to hibernate during winter in deep water on the bottom of continental-shelf slopes. This view has perpetuated in the literature even though the 'hibernation theory' has not been tested directly. We have now tracked basking sharks for the first time over seasonal scales (1.7 to 6.5 mo) using 'pop-up' satellite archival transmitters. We show that they do not hibernate during winter but instead undertake extensive horizontal (up to 3400 km) and vertical (> 750 m depth) movements to utilise productive continental-shelf and shelf-edge habitats during summer, autumn and winter. They travel long distances (390 to 460 km) to locate temporally discrete productivity 'hotspots' at shelf-break fronts, but at no time were prolonged movements into open-ocean regions away from shelf waters observed. Basking sharks have a very broad vertical diving range and can dive beyond the known range of planktivorous whales. Our study suggests this species can exploit shelf and slope-associated zooplankton communities in mesopelagic (200 to 1000 m) as well as epipelagic habitat (0 to 200 m).

Reduced pH sea water disrupts chemo-responsive behaviour in an intertidal crustacean

Chemoreception is a key activity by which many aquatic animals perceive their environment, and therefore abiotic disruptions to this process could have serious impacts on the survival and fitness of individuals, and on species interactions. Hermit crabs are subject to cyclical reductions in the pH of the water in the intertidal rock pools that they inhabit. Such reductions may be further exacerbated by ongoing ocean acidification and/or leakage of carbon dioxide from geological storage sites and coastal upwelling events. Here we test the chemo-sensory responses of the hermit crab Pagurus bernhardus (Linnaeus) to a food odour under reduced pH conditions (pHNBS = 6.80). Acidifying the odour had no effect on its attractiveness indicating no permanent degradation of the cue; however, the pH of the sea water did affect the crabs' responses. Hermit crabs kept and tested in reduced pH sea water had lower antennular flicking rates (the ‘sniffing’ response in decapods); were less successful in locating the odour source, and showed an overall decline in locomotory activity compared to those in untreated sea water. Analysis of their haemolymph revealed a greater concentration of chloride ions ([Cl−]) in the reduced pH treatment group, suggesting iono-regulatory disruption; however, there was no correlation between [Cl−] and locomotory activity, suggesting a specific effect on chemoreception. This study shows that the chemo-responsiveness of a crustacean may be influenced by both naturally occurring pH fluctuations and future anthropogenically-induced changes in ocean pH.

Scale-dependent foraging ecology of a marine top predator modelled using passive acoustic data

1.Understanding which environmental factors drive foraging preferences is critical for the development of effective management measures, but resource use patterns may emerge from processes that occur at different spatial and temporal scales. Direct observations of foraging are also especially challenging in marine predators, but passive acoustic techniques provide opportunities to study the behaviour of echolocating species over a range of scales. 2.We used an extensive passive acoustic data set to investigate the distribution and temporal dynamics of foraging in bottlenose dolphins using the Moray Firth (Scotland, UK). Echolocation buzzes were identified with a mixture model of detected echolocation inter-click intervals and used as a proxy of foraging activity. A robust modelling approach accounting for autocorrelation in the data was then used to evaluate which environmental factors were associated with the observed dynamics at two different spatial and temporal scales. 3.At a broad scale, foraging varied seasonally and was also affected by seabed slope and shelf-sea fronts. At a finer scale, we identified variation in seasonal use and local interactions with tidal processes. Foraging was best predicted at a daily scale, accounting for site specificity in the shape of the estimated relationships. 4.This study demonstrates how passive acoustic data can be used to understand foraging ecology in echolocating species and provides a robust analytical procedure for describing spatio-temporal patterns. Associations between foraging and environmental characteristics varied according to spatial and temporal scale, highlighting the need for a multi-scale approach. Our results indicate that dolphins respond to coarser scale temporal dynamics, but have a detailed understanding of finer-scale spatial distribution of resources.

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish. Behavioral Ecology and Sociobiology, 68, 1995-2003.

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.