Abundance, behaviour and habitat use patterns of Indo-Pacific bottlenose dolphins (\u3cem\u3eTursiops aduncus\u3c/em\u3e) in the Clarence and Richmond River estuaries in northern New South Wales, Australia

Bottlenose dolphins (Tursiops spp.) are a widely studied species in marine habitats, however, information on estuarine populations in Australia is very limited. To fully understand the importance of estuaries as habitats for dolphins there needs to be clear quantitative data on dolphin populations and their habitat use in estuaries. This study provides the first published data on Indo-Pacific bottlenose dolphins (Tursiops aduncus) abundance estimates, site fidelity, individual ranging patterns, habitat use, flood impacts and sexual segregation patterns over a 3- year period in the Clarence River (CR) and Richmond River (RR) estuaries in northern New South Wales, Australia. The results indicate that, at present, the CR estuary is sustaining a larger dolphin community with a predominantly resident population compared to the RR estuary, which supports a smaller community with lower site fidelity. The CR estuary dolphin abundance estimate of 71 (62-81 95% CI, CV = 0.07) is more than twice the size of the RR estuary abundance estimate of 34 (19- 49 95% CI, CV = 0.23). Differences in site fidelity were observed between the estuaries with 60% and 37% of identified dolphins determined as residents, 26% and 21% as occasional visitors, and 14% and 42% as transients for the CR and RR, respectively. Resource partitioning was apparent in both estuaries with the mean distance resident dolphins were found upstream from the River mouth being greater than for the occasional visitors and transients. Tursiops aduncus was seen all year round in the CR and RR estuaries, with peak sightings occurring in spring at both sites. In the CR the dolphin population showed consistent seasonal fluctuations, whereas this did not occur in the RR population. In the CR the largest spatial distribution of dolphins in the estuary was observed in spring and winter, while in summer they were primarily restricted to the main estuary channel. Different behaviours that were observed; feeding, socialising, travelling, and milling and resting, were found to be influenced by season, tidal phase and tidal range. In both estuaries the core habitat areas used by the population for feeding consisted of areas with considerable slope near the edge of tidal sand banks, adjacent to deeper channels at the entrances of canals, creeks or artificial breakwalls. In addition, the core habitat areas used by the population for milling and resting behaviour in both estuaries occurred in shallow, sheltered areas, often associated with seagrass beds. Socialising occurred more frequently in the CR throughout most of the deeper waters of the estuary, whereas in the RR it was primarily restricted to a small area of medium depth in the estuary. The major determinant of T. aduncus occupancy in the two estuaries was the flood events that occurred, which resulted in the dolphins abandoning the estuary. The mean predicted probabilities for sighting dolphins during non-flood periods were 0.87 and 0.71, during a flood 0.21 and 0.04, and during a post-flood recovery period 0.83 and 0.80 in the CR and RR, respectively. Principal Components Analysis (PCA) showed that when the dolphins were absent from the estuaries, three components were extracted from the water quality parameters in the CR, and two components in the RR. High loadings from the PCA were associated with the changing salinity, turbidity, pH, dissolved oxygen and temperature associated with the flood events. The return of the dolphins to the estuary following a flood depended on the length and severity of the flood event, but generally the dolphins seemed to prefer waters with salinity levels above 29 ‰. This could be associated with higher salinities being important for their physiological health, or because their prey returned to the estuaries during these higher salinity conditions, more likely a combination of both of these factors. Sexual segregation patterns were observed in T. aduncus populations with mixed gender and female groups mainly recorded in the CR, whereas in the RR female groups dominated the estuary. In the CR, significant differences occurred in sightings between the mixed and female groups in relation to water depth and behaviour. Mixed gender groups were sighted predominantly in deeper water and were involved in social behaviour including sexual behaviour and male herding of females. In contrast, the female groups were observed across all water depths, predominately feeding and also participating in more milling and resting behaviours. The high occurrence of aggressive herding behaviour by males in the CR was significantly different at varying depths, tides and seasons, occurring more often in deeper water, at higher tides and in non-breeding seasons. Female groups were found to utilise the small, shallow tributaries and travel for longer distances up these smaller tributaries than the mixed groups, which were concentrated in the deeper main channel of the estuary. The deeper water of the channels may facilitate the males in herding the females, while the female groups’ habitat selection of shallow estuary areas may provide a sanctuary from aggressive males, access to suitable prey items or prey density for mothers and their calves, or a combination of these factors. This study has provided the first detailed research on T. aduncus dolphin population dynamics, habitat use, occupancy and sexual segregation patterns in two Australian subtropical estuaries. To ensure the long-term survival of both of these dolphin populations, management of future increased anthropogenic disturbances from boat traffic, pollution, dolphin watching, industrial or urban development, over-fishing and habitat degradation of the catchment is needed. Good quality water conditions, the protection of the core feeding areas and small shallow tributaries for females and their calves, and sheltered areas for resting behaviour all need to be maintained for the continued conservation of these important dolphin populations

Seasonal variation and tidal influences on estuarine use by bottlenose dolphins (Tursiops aduncus)

In order to show that dolphins use estuary habitats differently depending on the season and tidal state, possibly in response to prey distribution, temperature, risk of stranding and accessibility, Indo-Pacific bottlenose dolphins (Tursiops aduncus) were observed year-round during a 3-year study in the Clarence River estuary (CR) and Richmond River estuary (RR) in northern New South Wales, Australia. Peak dolphin sightings occurred during the spring season and one or 2 h prior to high tide. The spatial distribution of the dolphins in each estuary was analysed using the distance in kilometres that the dolphins travelled upstream with seasons and tidal phase as determinants. A General Linear Model showed that in the CR the dolphin spatial distribution in the estuary was not determined by season (F = 0.434, df = 3, P = 0.729) but was by tidal phase (F = 9.943, df = 3, P \u3c 0.001) and the interaction between season and tidal phase (F = 3.398, df = 9, P \u3c 0.002). However, in the RR the spatial distribution of the dolphin use of the estuary was not determined by either season (F = 1.647, df = 3, P = 0.194) or tidal phase (F = 0.302, df = 3, P = 0.824). In the CR, the spatial distribution of the dolphins was largest on high and flood tides. This pattern of spatial distribution may occur because the CR is a relatively shallow estuary and this increased spatial distribution may reflect a lower stranding risk and an increase in accessibility of shallow areas during periods of higher tide. These areas could also provide access to their preferred prey items of sea mullet (Mugil cephalus) and sand whiting (Sillago ciliata)

Mixed-sex groups' kernel density estimates (KDE) displaying aggressive coercive behaviour in the Clarence River estuary (N = 336).

<p>Mixed-sex groups' kernel density estimates (KDE) displaying aggressive coercive behaviour in the Clarence River estuary (N = 336).</p

Percentage of dolphin encounters in varying water depths for female and mixed-sex groups in the Clarence River estuary per season.

<p>The sum for the four seasons amounts to 100% for each water depth. Seasons are in order Spring, Summer, Autumn and Winter.</p

Frequency of dolphin behaviours for female (a), male (b), and mixed-sex (c) groups in the Clarence River estuary.

<p>* 76% of the total social interaction time (N = 442) of mixed-sex groups involved aggressive coercive behaviour.</p

Mixed-sex groups' kernel density estimates (KDE) for all behaviours from the Clarence River estuary (N = 1118).

<p>Mixed-sex groups' kernel density estimates (KDE) for all behaviours from the Clarence River estuary (N = 1118).</p

Boat-survey effort and direct dolphin observation (DDO) periods during each season.

<p>Boat-survey effort and direct dolphin observation (DDO) periods during each season.</p

Female groups' kernel density estimator (KDE) for all behaviours from the Clarence River estuary (N = 539).

<p>Female groups' kernel density estimator (KDE) for all behaviours from the Clarence River estuary (N = 539).</p

Map of the study sites in northern New South Wales, Australia at the top and a detailed map of the Clarence River estuaries indicating the survey areas.

<p>Map of the study sites in northern New South Wales, Australia at the top and a detailed map of the Clarence River estuaries indicating the survey areas.</p