POPULATION DYNAMICS AND SITE FIDELITY OF BOTTLENOSE DOLPHINS (TURSIOPS ADUNCUS) IN TWO ESTUARIES IN SUBTROPICAL NORTHERN NEW SOUTH WALES, AUSTRALIA
CHRISTINE A. FURY*
Southern Cross University Whale Research Centre
School of Environmental Science and Management
Southern Cross University
PO Box 157, Lismore, NSW, Australia 2480
* Email email@example.com
Telephone +61 (02) 6620-3815
Fax +61 (02) 6621-2669
Population dynamics, occupancy and residency patterns were examined over a 3-year period for Indo-Pacific bottlenose dolphin (Tursiops aduncus) populations in the Richmond River (RR) and Clarence River (CR) estuaries in northern New South Wales, Australia. Mark-recapture analyses estimated a RR population size of 41 (95% CI 32-72) and a CR population size of 67 (95% CI 66-80). High site fidelity was observed in both estuaries with 61% and 77% of identified dolphins determined as long-term residents, 17% and 13% as intermittent residents, and 22% and 10% as transients for RR and CR, respectively. The populations in the RR and CR are separate. High numbers of calves and neonates were recorded from the total number of identified dolphins in the RR (21.2%) and high numbers of juveniles occurred in the CR (22.4%).
Bottlenose dolphins (Tursiopsspp.) are cosmopolitan species found throughout tropical and temperate seas and oceans worldwide. They inhabit a large range of marine habitats including deep oceans to shallow coastal regions, inshore lagoons and estuaries (Connor et al., 2000). However, insufficient information is available about the status of Bottlenose dolphin populations and there is no published information on bottlenose populations in Australian estuaries. Estimated population size of bottlenose dolphins inhabiting estuaries in other locations range from 40 from the Sado estuary, Portugal (dos Santos and Lacerda, 1987) to 129 in the Moray Firth, Scotland (Wilson et al., 1999b).
The social ecology of bottlenose dolphins varies in relation to different habitats. In more protected coastal and estuarine environments bottlenose dolphin populations usually associate in small groups, show higher site fidelity, feed primarily on dispersed prey and belong to relatively small local populations (Bryden et al. 1998; Defran and Weller 1999). Conversely, in more open habitats such as pelagic waters, bottlenose dolphins will usually associate in large groups or aggregations, show low site fidelity, and feed on large prey patches that are distributed in clumps, separated by vast expanses of water (Bryden et al., 1998, Defran and Weller, 1999, Würsig, 1978, Shane et al., 1986).
The importance of estuaries as an area of habitat choice for bottlenose dolphins needs to be understood. Baseline data on their population size and occupancy is essential for their effective conservation and management (Shirakihara et al., 2002). This is particularly important in Australian estuaries where human populations are rapidly expanding and various direct and indirect threats to dolphins will increase. These threats include boat traffic (Constantine, 2001, Bejder et al., 2006, Kelly et al., 2004, Dunn et al., 2001, Allen and Read, 2000), recreational and commercial fishing (Bearzi et al., 2006, Charlton et al., 2006, Vidal, 1993), pollution (Wells et al., 2005, Wilson et al., 1999a), and habitat degradation (Bearzi et al., 1997).
The aim of this 3-year photo-identification, mark-recapture study was to determine the population size, occupancy and residency of Indo-Pacific bottlenose dolphins (T. aduncus) in two Australian estuaries in northern New South Wales, Australia.
Study sites – Richmond and Clarence rivers
The Richmond River and the Clarence River flow through adjacent river valleys in northern New South Wales (NSW), Australia, and these estuaries and separated by 65km (Fig. 1). The Richmond River is classified as a wave-dominated delta (Ryan et al., 2003) located on the subtropical north coast of New South Wales, Australia (153°35'E 28°53'S). It has a catchment area of approximately 6,861 km2 , a waterway area of 14.3 km2 and volume of 67.7 m³ × 106 with a mean depth of 4.7 m (Walsh et al., 2004, Eyre, 2000). The estuary is well mixed even in the dry season, with a maximum tidal influence of 60 km from the mouth and semidiurnal tides with amplitudes that range from 0.5-1.5 m for neap tides and 0.2-2 m for spring tides (Hossain et al., 2001).
The Clarence River is classified as a wave-dominated estuary (Ryan et al., 2003) located on the north coast of New South Wales, Australia (153°21'E 28°26'S) (Fig 1). It is the largest coastal river in northern New South Wales (West, 2002), and has a catchment area of approximately 22,446 km2 and a waterway area of 89 km2, mean depth of 2.3 m, volume of 204.7 m³ × 106 and a mean annual rainfall of 1075 mm. The approximate length of the river tidal influence is about 60 km from the mouth. The mean spring tidal range at the entrance is 1.34 m (Walsh et al., 2004, Eyre, 2000).
Fig. 1 – Map of the study sites in northern New South Wales, Australia (1) and a detailed map of both estuaries of the Richmond River (2) and Clarence River (3) indicating the survey areas (shaded), coast line (line) and land survey observational points (triangles).
Boat-based surveys were completed in the Richmond River (RR) and Clarence River (CR) over 3 years from October 2003 to September 2006. Surveys were done predominantly using a 3.8m aluminium boat powered by a 15hp 2-stroke engine, however, for the first year a 4.2m aluminium boat powered by an 8hp 2-stroke engine was mainly used in the CR. During the surveys the boat was kept at a steady speed of ~6 knots and 1-3 observers maintained a constant visual search for dolphins. All surveys were done in calm conditions (i.e., Beaufort sea state 3) in daylight hours between 0600 and 1800 h. The route for each survey was random and the entire study area was surveyed until a dolphin group was located (Fig. 1). Surveys ranged over a ~22km area in the RR and ~ 28 km area in CR upstream from the river mouth. Different times of the day, tidal periods, weather conditions, months and all seasons were used for surveys to minimise bias.
Photo-identification and analysis
Photographic identification of bottlenose dolphins relies on matching unique marks and nicks on dorsal fins and flanks of individuals dolphins (Würsig and Jefferson, 1990, Würsig and Würsig, 1979, Wilson et al., 1999b). The trailing-edge of the dorsal fin, in particular, is easily damaged resulting in a unique dorsal fin profile for each dolphin. Focal follows of dolphin groups were done during the surveys where a focal individual or group were followed to the maximum possible extent up to 3 hours during each sample period. Dolphin groups were followed from a distance of up to 50m, but mostly from ~ 30m which was the approved limit under the research permits.
Photographs were taken predominantly using a Canon EOS D20 digital camera and Canon zoom lens EF 70-200mm f/2.8G IS USM Image Stabilizer. On some occasions, photographs were taken using a Canon EOS D10 camera or a Nikon D100 camera with a Nikon lens VR 70-200mm f/2.8G with a Nikon AF-S 1.4x teleconverter.
Upon encountering a group of dolphins the aim was to photograph each dolphin’s dorsal fin from both sides. The group size encountered during boat surveys ranged from 1-34, however, was most often between 2-5 dolphins; hence, photographing all dolphins in a group was usually achievable. Each photograph was categorised as fair, good or excellent and only the good and excellent photographs of each dolphin were compared to the catalogued photographs using Adobe Photoshop version 6.0 software (Adobe Systems, San Jose, CA) or Nikon browser (version 6).
The program MARK (White and Burnham, 1999) was used to estimate the population sizes of bottlenose dolphins in the two study areas. Photo-identification data from the 3 years were divided into seasons with each season per year creating a capture occasion. The monthly surveys were usually completed within 5 consecutive days per month, particularly in the Clarence River. Consequently, some dolphins may have been missed during a particular month, because surveys on consecutive days occurred during periods of similar tides, times or weather conditions. Therefore, seasonal data rather than monthly data were used to avoid missing individual dolphins that were not sighted during a particular month.
To determine if the populations were open or closed (to births, deaths, immigration, emigration) the discovery curve was plotted to analyse the cumulative number of marked individuals. Data from the discovery curve levelled off towards the end of the study period, strongly suggesting that dolphin populations in both the RR and CR were closed (Williams et al., 1993). The models for mark-recapture analysis in MARK for a closed population include Mt (accounting for time), Mh (accounting for heterogeneity of probability of capture), and Mth (both of time and heterogeneity). Although the term Mb accounting for behavioural difference was initially included, because behavioural responses are not known to occur from non-invasive photo-identification surveys, this term was subsequently disregarded (after Wilson et al. 1999a). To comply with the assumptions for mark loss during the study for closed capture analysis, only dolphins with distinctive and permanent marks were used, only good and excellent category photographs were included, and only one experienced person (CF) was responsible for cataloguing and recognising dolphins to minimise bias. The final population models used for both study sites accounted for heterogeneity of capture probability because the total home range of all the dolphins was not known since surveys did not include the area outside of the estuaries where the dolphins are known to occur. Calves were also excluded from the analyses, as their probability of capture was not exclusive from that of their mothers (Wells and Scott, 1990).
To calculate the proportion of unmarked individuals (non-calves) in the population, the total number of unmarked individuals was divided by the total number of individual dolphins identified after Stensland et al. (2006). To calculate the proportion of calves in the population, the total number of identified calves was divided by the total number of marked dolphins. The population estimates and confidence intervals were then adjusted to compensate for the number of unmarked individuals (non-calves) in the population after Stensland et al. (2006) and to include calves in the population estimation.
Boat-based survey data were combined with land-based survey data that was completed concurrently during the 3 years of the study to more accurately determine the dolphins’ residency patterns and resight rate in the study area. Land-based surveys were done from October 2003 to September 2006 in the RR and CR on a seasonal basis (~4 per season) for a period of 3 years. Not all dolphins could be photographed during land surveys because of their distance in the estuary from the observation sites (Fig. 1). The land-based survey data were only used for the 71 marked dolphins sighted in the estuaries during the boat-based surveys.
To determine residency patterns, definitions were adapted from those of Zolman (2002), which are based on presence or absence of individually identified dolphins per season. Each survey year was divided into four seasons: spring (September-November), summer (December-February), autumn (March-May) and winter (June-August) for the 3 years, resulting in 12 seasonal periods in total.
Dolphins were defined as long-term residents when they were sighted in at least 3 of the 4 seasons, regardless of year, and with sightings in a minimum of 6 of the 12 seasonal survey periods in the study area. Intermittent residents were dolphins that were sighted in 2-5 of the 12 seasonal survey periods, with no sightings in consecutive seasons. Transients were dolphins sighted in only one season and/or in consecutive seasons, but not again in the study area, during the study period. Calves with known mothers were classified in the same category as the mother while still dependant.
Group size and composition
A group of dolphins was defined as all dolphins within a 100m radius of each other and engaged in similar activities (Lusseau et al., 2005, Wells et al., 1987, Wilson, 1995, Irvine et al., 1981). The definitions of dolphin age classes followed those of Bearzi et al. (1997) and comprised of neonate, calf, juvenile and adult categories. Mothers were defined as adult dolphins seen in repeated, close association with a calf or neonate on at least two independent occasions. The calf was often seen swimming in the infant position (Mann and Smuts, 1999). Data were excluded from analysis if group composition could not be established.
All analyses, apart from population estimation, were done using SPSS for Windows (version 11.5). To confirm that there was equal survey effort across all seasons at each site, the survey hours per season were compared using a Pearson Chi-square test. To determine if there were any differences in residency patterns for the marked individuals including calves, a Pearson Chi-square test was used to compare percentages of long-term residents, intermittent residents, and transients. The group sizes were then analysed for differences between seasons and among the 3 years of the study with a non-parametric, independent sample, Kruskal-Wallis ANOVA. A non-parametric Mann-Whitney U test was done to determine if there was a difference between the mean dolphin group size between groups containing calves and those without calves.
Survey and Photo-identification effort
Boat-based surveys were done for 3 years from October 2003 to September 2006 in the RR and CR on a seasonal basis (4-12 surveys per season). A total of 100 and 87 boat-based surveys were completed during the 3-year study comprising of 993.5 hours (RR 525; CR 468.5), with 213 and 541 dolphins sighted in the RR and CR, respectively. The median survey duration was 360 mins (range 120-510 mins). Survey effort was similar across all seasons in both sites: Spring 124.5h and 111.5h, Summer 124h and 121.5h, Autumn 131h and 124.5h, and Winter 146h and 111h in the RR and CC, respectively (RR: χ² = 2.35, df = 3, P = 0.5; CR: χ² = 1.27, df = 3, P = 0.74).
A total of 120 land-based surveys were completed in the RR and 128 surveys in the CR over 3 years comprising of 722 hours (RR 354; CR 368) with 965 (RR) and 1,420 (CR) dolphin sightings. All survey durations were 3 hours, with 0-7 surveys per month and 15 hrs – 39 hrs (RR) and 24 hrs - 36 hrs (CR) per season (mean and median = 4).
During the boat-based surveys from October 2003-September 2006 photographs of the 747 dolphins encountered resulted in more than 7,400 photographs. At the conclusion of the research, the total catalogued numbers of identifiable dolphins, including calves, was 23 in the RR and 48 in the CR.
Dolphin groups were encountered on 55% (n = 100) and 76% (n = 87) of all surveys in the RR and CR, respectively. Multiple dolphin groups were observed on 24% (RR, N = 87) and 24% (CR, N = 142) of individual surveys. In the RR, 18% of surveys (n = 16) contained two dolphin groups and 6% (n = 5) of surveys had three to five dolphin groups present. In the CR, two dolphin groups were observed in 13% (n = 19) of the surveys, 5% (n = 7) had three dolphin groups and 6% (n = 8) had four to nine dolphin groups present.
Discovery rate and Mark rate
The cumulative discovery rate of uniquely marked dolphins was plotted to determine if the population was open or closed (Fig. 2). The dolphin sightings were expressed by seasons and after the first year 61% (RR) and 74% (CR) of the total dolphins had been identified, and 91% (RR) and 90% (CR) had been identified by the end of the second year. All dolphins were identified by spring 2005 (RR) and summer 2005/6 (CR). Therefore, the population was considered to be closed for mark-recapture analyses (Williams et al., 1993). The percentage of dolphins with unique marks in the RR population was 85.5% and in the CR population was 78.3%.
Fig. 2 - Discovery curves of the cumulative number of bottlenose dolphins identified from permanent marks per season between October 2003 and September 2006 in the Richmond River and Clarence Rivers, northern New South Wales.
The population estimates were derived from the program MARK to determine the models best suited to each population (Table 1). There was no interchange between dolphins found in each estuary, therefore, they were assumed to be separate populations. The models selected for each population were determined by ranking them according to their criterion values (maximum = 1). These were Mh for the Richmond River population and Mth for the CR population. The adjusted population estimates including unmarked dolphins and calves were 41 (95% CI 32-72) for the RR, and 67 (95% CI 66-80) for the CR (Table 1).
Source of variation
Table 1 - Adjusted estimates for total bottlenose dolphin population sizes. The model selected for each population by the high criterion value is in bold and underlined.
Resight rate and Residency patterns
There was considerable variation in the resighting rate of the 71 (RR = 23, CR = 48) marked individuals observed in the two study sites from the boat and land-based surveys. Some dolphins were resighted up to 37 times in total and up to 10 times in a single season in the RR, and up to 33 times in total and up to 8 times in a single season in the CR (RR: mean 13, median 9; CR: mean 16.6, median 15). In contrast, a few individuals were only seen once at each site during the boat and land-based surveys (RR: 22% N = 5; CR: 6% N = 3).
Residency patterns for the marked individuals, including calves, indicated that 61% (n = 14) and 77% (n = 38) were long-term residents, 17% (n = 4) and 13% (n = 6) were intermittent residents, and 22% (n = 5) and 10% (n = 6) were transients in the RR and CR, respectively. These proportions were significantly different with respect to residency patterns at both sites (RR: χ² = 34.8, df = 2, p = 0; CR: χ² = 85.9, df = 2, p = 0).
No intermittent residents were only seen during one particular season during the 3 years. All intermittent residents had a gap in sightings at some stage in the 12 seasonal sampling periods. Transient dolphins in the RR generally appeared in spring (88%), but in the CR only 43% of the transients were sighted in spring, and the other individuals were sighted throughout the other seasons.
Group size and composition
The mean dolphin group size was 3.1 (SE 0.22, interquartile range = 3, Median = 2) for the RR population and 4.6 (SE 0.28, interquartile range = 4, median = 4) for the CR population (Fig. 3). There were no significant differences in the mean group sizes in different seasons at both sites (ANOVA: RR: z = 3.147, df = 3, p = 0.369; CR: z = 1.708, df = 3, p = 0.635) or among years (RR: z = 0.341, df = 2, p = 0.843; CR: z = 0.895, df = 2, p = 0.639).
Age category data of 985 dolphin sightings (RR 222, CR 708) from boat surveys indicated that 66.4% and 70% were adults, 14% and 22.4% were juveniles, and 21.2% and 7.6% were calves and neonates in the RR and CR, respectively. The mean sizes of dolphin groups containing calves were 4.2 (RR) and 6.4 (CR), which was significantly larger than the mean sizes of dolphin groups without calves, which were 2.2 (RR) and 3.9 (CR) at the two sites (RR: Mann-Whitney, U = 507.5, p = 0; CR: U = 1,693, p = 0).
Fig. 3 – Frequency of dolphin group sizes sighted from boat surveys from October 2003-September 2006 in the Richmond River (n = 100) and Clarence Rivers (n = 87).
Mother dolphin groups
In the RR there were 17 marked adults, 4 of which were mothers and all 4 were long-term residents (100%). In contrast, in the CR there were 34 marked adults with 10 classified as mothers. Of those mothers, 70% were long-term residents (N = 7), 20% were intermittent residents (N = 2), and 10% (N = 1) were transients (χ² = 62, df = 2, p = 0).
This study has provided the first published estimates of population sizes and residency patterns for bottlenose dolphins inhabiting estuaries in Australia. The Richmond River estimated population size of 41 and the Clarence River estimated population size of 67 are both moderate population sizes relative to the size of these estuaries. Published population data are available for bottlenose dolphins inhabiting estuaries in a few other locations worldwide (Table 2). The population size of 40 in a survey area of approximately 150km² in the Sado estuary, Portugal (dos Santos and Lacerda, 1987, Harzen, 1998) is smaller than both the Richmond (14.3 km²) and Clarence (89 km²) estuary populations. The Sado estuary has industrial pollution and over-fishing problems that are thought to contribute to the low dolphin numbers (dos Santos and Lacerda, 1987). In contrast, the Moray Firth bottlenose dolphin population was estimated to be 129, which reflects the large habitat available within this large open estuary (Wilson et al., 1999b).
The Richmond River and Clarence River dolphin populations’ exhibit high site fidelity, with 61% and 77%, respectively, determined as long-term residents. These estuaries represent only part of a likely larger home range for these dolphins, as they were sighted entering and exiting the river mouth as well as proceeding further upstream beyond the survey areas. Other estuarine bottlenose dolphin populations show variations in residency patterns. Some populations are similar to the RR and CR, with a predominant resident population, which has been reported from the Moray Firth (Wilson et al., 1997) and Southern Brazil Laguna’s canal and Imarui-Santo Antonio’s lagoon (Simoes Lopes and Fabian, 1999), and the Sado estuary in Portugal (dos Santos and Lacerda, 1987, Harzen, 1998). In contrast, a large transient dolphin population dominates Calibogue Sound in South Carolina (Gubbins, 2002a), an open estuarine system in Cedar Keys, Florida (Quintana-Rizzo and Wells, 2001) and the Stono River estuary, South Carolina, USA (Zolman, 2002).
The percentage of calves and neonates in the RR population was high at 21.2% compared to other study sites such as Jervis Bay, NSW, Australia with 5.9% (Möller and Harcourt, 1998), Florida, USA with 8.2% (Irvine et al., 1981), and in the northern Adriatic Sea where only 8.2% juveniles, and 6.6% calves and newborns were recorded in the population (Bearzi et al. 1997). The Clarence River had a slightly higher percentage of calves and neonates (7.6%), and a high percentage of juveniles (22.4%) compared to the northern Adriatic Sea population and the RR (14%).
Most of the mothers in the RR and CR were residents or intermittent residents, and 29% of adult dolphins in the RR were mothers. Bottlenose dolphins in the Stono River estuary showed a similar pattern with mothers in the resident population making up one-third of the total population. The CR had 76% resident dolphins of which 18% were current mothers.
Do estuaries provide an important nursery habitat for mothers and their calves in some populations? Mothers with neonates tend to spend more time in habitats that offer protection, preferring shallow waters with high prey availability (Wang et al., 1994, Scott et al., 1990, Ballance, 1992, Zolman, 2002). The main dolphin group in the Richmond River consists of a group of females with their calves, which may reflect habitat selection by lactating females who reside in estuaries because of their higher caloric needs (Whitehead and Mann, 2000).
Calibogue Sound & surrounds including coastal area, South Carolina, USA
478 individually identified
Richmond River, Australia
41 (95% CI = 31-70)
Clarence River, Australia
67 (95% CI = 67-84)
Table 2 - Table of published bottlenose dolphin population sizes and estuary sizes worldwide.
Small animal populations are more prone to extinction (McCarthy and Thompson, 2001), particularly for long-lived animals with a small reproductive output such as bottlenose dolphins (Wells, 1991). Additional land-based survey data (CF, unpublished data) indicate that a substantial number of bottlenose dolphins occur outside the Richmond River estuary, whereas few dolphins were observed outside of the Clarence River estuary. These trends were reflected in the numbers of transient dolphins entering these estuaries, with 22% in the RR and 10% in the CR. All transients in the RR were sighted in spring, and were sighted mostly socialising with the resident dolphins and are therefore likely to be males in search of females for breeding. In contrast, in the CR transients were sighted in most seasons and 40% were sighted singularly and feeding (CF, unpublished data).
Genetic data also supports the premise that the coastal dolphin population outside of the Clarence River is small, and preliminary analysis has shown the population to be genetically distinct (J. Wiszniewski and L. Moller, pers. comm.). Therefore, less genetic exchange is occurring in this population which makes the CR population more vulnerable to local extinction. If events such as disease outbreak or reduction in fish stocks occur to reduce the population size then the CR dolphins will have less conspecifics in the region to re-stock the population (Thompson et al., 2000, McCarthy and Thompson, 2001).
The Richmond River and Clarence River regions are likely to face a 34% and 30% increase in their human populations within the next 30 years, respectively (Transport and Population Data Centre, 2004). Increasing human populations in coastal and river catchments require improved management practices around the coasts and rivers to reduce pressures on vulnerable dolphin populations. A dam is proposed for the Clarence River which would reduce the river flow and potentially decrease fish stocks in the estuary that supports the dolphin population (Paugy et al., 1999).
The results of this study indicate that, at present, the Richmond River and Clarence River estuaries are sustaining a predominantly resident dolphin population with relatively high proportions of mothers, calves and juveniles in these sheltered nursery habitats. However, to ensure the long-term survival of these dolphin populations, management of future increased anthropogenic disturbances from boat traffic, pollution, over-fishing and habitat degradation is essential.
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