Wildlife Research, 2012,24,185-208

Distribution and Abundance of Dugongs, Turtles,

A. R. PreenA, H. MarshA, I. R. LawlerA, R. I. T. PrinceB and R. ShepherdC

ADepartment of Tropical Environment Studies and Geography, James Cook University

WA, 6065, Australia.

Abstract
Strip-transect aerial surveys of Shark Bay, Ningaloo Reef and Exmouth Gulf were conducted
during the winters of 1989 and 1994. These surveys were designed primarily to estimate the
abundance and distribution of dugongs, although they also allowed sea turtles and dolphins, and,
to a lesser extent, whales, manta rays and whale sharks to be surveyed. Shark Bay contains a
large population of dugongs that is of international significance. Estimates of approximately 10000
dugongs resulted from both surveys. The density of dugongs is the highest recorded in Australia
and the Middle East, where these surveys have been conducted. Exmouth Gulf and Ningaloo Reef
are also important dugong habitats, each supporting in the order of 1000 dugongs. The estimated
number of turtles in Shark Bay is comparable to the number in Exmouth Gulf plus Ningaloo Reef
(7000-9000). The density of turtles in Ningaloo Reef and, to a lesser extent, Exmouth Gulf is
exceptionally high compared with most other areas that have been surveyed by the same
technique. Shark Bay supports a substantial population of bottlenose dolphins (2000-3000
minimum estimate). Exmouth Gulf and Ningaloo Reef were not significant habitats for dolphins
during the winter surveys. Substantial numbers of whales (primarily humpbacks) and manta rays
occur in northern and western Shark Bay in winter. Ningaloo Reef is an important area for whale
sharks and manta rays in autumn and winter. The Shark Bay Marine Park excludes much of the
winter habitats of the large vertebrate fauna of Shark Bay. In 1989 and 1994, more than half of all
the dugongs were seen outside the Marine Park (57á4 and 50á7%, respectively). Approximately
one-third to one-half of turtles and dolphins were seen outside the Marine Park (in 1989 and 1994
respectively: turtles, 43 and 27%; dolphins, 47 and 32%). Almost all the whales and most of the
manta rays were seen outside the Marine Park. Expansion of the Shark Bay Marine Park, to bring it
into alignment with the marine section of the Shark Bay World Heritage Area, would facilitate the
appropriate management of these populations. This would also simplify the State - Commonwealth
collaboration necessary to meet the obligations of World Heritage listing. The coastal waters of
Western Australia north of the surveyed area (over 6000 km of coastline) are relatively poorly
known and surveys of their marine megafauna are required for wise planning and management.

Introduction
Western Australia has an extensive coastline, extending 12500 km over 21o of latitude and 16o
of longitude. Southern and western coastal waters are relatively well known (Wilson et al. 1979;
Pearce and Walker 1991), but knowledge of the habitats and marine fauna decreases northwards
from the populated south-west (Marine Parks and Reserves Selection Working Group 1994). The
central western coast, between 22 and 27¡S, includes a number of areas of high conservation
status, including the Shark Bay World Heritage Area and Ningaloo Marine Park. Effective
management of these areas requires sound knowledge of their environments, including their
importance for threatened and high-profile species. Hence, the distribution and abundance of

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186 A. R. Preen et al

dugongs and other megafauna of this region was assessed by aerial survey during the winter of
1989. Megafauna, in this context, are large marine vertebrates that can be surveyed from the air.
The 1989 survey established Shark Bay as a site of particular significance for dugongs (Marsh
et al. 1994). It also documented a distribution pattern that was at odds with expectations (Anderson
1982, 1986; Marsh et al. 1994). Given the management implications of these findings, and the
need to determine whether the population was stable, increasing or decreasing, a second regional
survey was conducted in 1994. Five years is the recommended interval between repeat surveys of
dugongs to detect population trends (Marsh and Saalfeld 1989a). Such surveys have demonstrated
population stability in some areas (northern Great Barrier Reef) and alarming declines in others
(central and southern Great Barrier Reef; Marsh et al. 1996). Here we document and compare the
distribution and abundance of dugongs, turtles and dolphins in Shark Bay, Ningaloo Reef and
Exmouth Gulf during the winters of 1989 and 1994. We also provide information on the distribution
of whales, manta rays and whale sharks. Data on dugongs from the 1989 survey of Shark Bay
have been presented previously (Marsh et al. 1994).
Shark Bay supports a dugong population of international significance, a large turtle population,
and is important for small and large cetaceans. While the Shark Bay World Heritage Area
encompasses virtually all the important non-migration habitat for these species in this region, the
Shark Bay Marine Park does not. This anomaly has the potential for confusion and conflict due to
the different management regimes that may be expected by State and Federal agencies in the
World Heritage Area outside the Marine Park.
Ningaloo Reef and Exmouth Gulf support important populations of turtles and are significant
habitats for dugongs. Manta rays and whale sharks are relatively common off the Ningaloo Reef in
winter. Ningaloo Reef is well protected in the Ningaloo Marine Park, but Exmouth Gulf, which
sustains an important trawl fishery and is covered by petroleum exploration permits, currently has
no special conservation status.

Methods
Study Areas
Shark Bay is a 13000-km2 embayment complex on the central Western Australian coast (25o
30’S, 113o30’E). It experiences a semi-arid to arid climate and precipitation (200Ð400 mm per
year) is greatly exceeded by evaporation (2000-3000 mm per year). Consequently, enclosed parts
of the bay, particularly Hamelin Pool, are hypersaline (>42%0). About half the bay is metahaline
(38Ð42%0), while the remainder, primarily the western and northern areas, has oceanic salinities
(35Ð40%0; Fig. 1) (Logan and Cebulski 1970).
Shark Bay has been inscribed on the World Heritage List in recognition of its outstanding
universal natural values (Department of Arts, Sport, the Environment, Tourism and Territories
1990; Department of Environment, Sport and Territories 1995). Seagrass meadows cover more
than 4000 km2 and are reported to be the largest in the world (Walker 1989). These meadows are
composed predominantly of the temperate species Amphibolis antarctica, although there is recent
evidence of large deep-water meadows dominated by the tropical species Halophila spinulosa
(Anderson 1994). The bay supports a population of approximately 10000 dugongs (Dugong
dugon), making it a habitat of international significance for this species (Marsh et al. 1994). Nesting
beaches at the northern tip of Dirk Hartog Island are used annually by as many as 800Ð1000
female loggerhead turtles (Caretta caretta), which may represent 70% of the Western Australian
breeding population (Prince 1994a). Monkey Mia, on the eastern coast of the Peron Peninsula, is
renowned for the contact between humans and some members of the local population of
bottlenose dolphins (Tursiops truncatus) (Smolker et al. 1992). The World Heritage Area includes
the entire bay complex, excluding nearshore waters around Carnarvon (Fig. 1). The southern and
eastern areas of the bay are included in the Shark Bay Marine Park and the Hamelin Pool Marine
Nature Reserve (Department of Conservation and Land Management 1994) (Fig. 1).
Ningaloo Reef (22o30’S, 113o48’E) is the largest fringing barrier reef in Australia and extends
260 km north-south along the western shore of the Cape Range Peninsula (Department of
Conservation and Land Management 1989) (Fig. 1). It is mostly enclosed by Ningaloo Marine Park
(Marine Parks and Reserves Selection Working Group 1994). Unlike most other coral reef systems
187 Megafauna of Coastal Waters off Western Australia

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188 Megafauna of Coastal Waters off Western Australia

in Australia, NingalooÕs near-shore reefs enclose a narrow lagoon varying in width from 200 m to
6 km and with an average depth of only 4 m (Department of Conservation and Land Management
1989). Because of the aridity of the hinterland, and consequent low run-off, the lagoon waters are
particularly clear (May et al. 1983), especially when compared with most inshore reef waters in
Australia. Ningaloo Reef is contiguous to Exmouth Gulf (22¡09S, 114¡249E), which lies on the
eastern side of the Cape Range Peninsula (Fig. 1). Exmouth Gulf is a relatively turbid environment,
characterised by areas of fringing mangroves, mudflats, rock pavements and softbottom habitats.
Early shoreline aerial surveys indicated that Exmouth Gulf was an important habitat for dugongs
(Prince et al. 1981).

Aerial Surveys
We conducted strip-transect aerial surveys of Shark Bay, Ningaloo Reef and Exmouth Gulf
between 4 and 13 July 1989 and between 21 and 30 June 1994. The surveys were conducted
during winter to take advantage of the relatively light winds, and hence better observation
conditions. The Shark Bay surveys encompassed the entire World Heritage Area, except for the
western shorelines of the barrier islands (Fig. 2). The Exmouth surveys covered all of Exmouth
Gulf, as defined by an eastÐwest line through the tip of North West Cape (Fig. 2). The Ningaloo
surveys encompassed only the near-shore waters of Ningaloo Marine Park (most of the park is
deep and oceanic and was excluded). Unsuitable weather prevented the survey of the southern
half of the Marine Park in 1989.
The methods followed those of Marsh and Sinclair (1989a, 1989b) and are detailed for the 1989
Shark Bay survey in Marsh et al. (1994). To calculate regional densities of fauna, Shark Bay was
divided into eight survey blocks (Fig. 2). Ningaloo Reef and Exmouth Gulf were treated as single
blocks, with a small
transects were 4á65 km apart (2á59 latitude), resulting in a coverage of 8á9Ð9á9%, except in
Shark Bay Blocks 0 and 2, where transects were 9á3 km apart, and coverage was 4á1%. The
same transects were flown on each survey, except for Ningaloo Reef. In 1989, 19 transects,
covering only the northern half of Ningaloo Marine Park, were surveyed (Fig. 2). In 1994, 43
transects (including the original 19), covering the full length of Ningaloo Marine Park, were flown.
The 1994 transects extended approximately 200 m seaward of the fringing reef, while the 1989
transects extended 0á8Ð4á5 km (mean = 2á3 km) further.
Surveys were conducted only under good weather conditions (Beaufort sea state <3; Table 1),
and we avoided flying during periods of severe glare (early morning, late afternoon and midday).
Sightings were recorded from a 200-m-wide strip on each side of the aircraft, from an altitude of
137 m, at an aircraft speed of about 185 km hÐ1.

Two isolated, independent observers were used on each side of the aircraft. From a
markÐrecapture analysis of sightings, perception-bias correction factors were derived to adjust the
results to allow for the animals visible, but missed by observers (Marsh and Saalfeld 1989a) (Table
2). Separate perception-bias correction factors were calculated for each side of the aircraft for
dugongs and turtles for Shark Bay and for the combined NingalooÐExmouth blocks. Owing to the
low number of dolphins recorded in Exmouth Gulf, the Shark Bay correction factor was used.
Insufficient dolphins were seen in Ningaloo to derive population estimates, so no correction factors
were needed.
Availability correction factors were derived to correct for the number of animals not at the
surface, and hence less likely to be available to observers, at the time the plane passed over
(Marsh and Sinclair 1989b) (Table 2). For dugongs, the proportion of sightings at the surface was
compared to the proportion at the surface in Moreton Bay, Queensland, where all dugongs feeding
in 2Ð3 m of water were visible. That proportion was determined from vertical aerial photographs.
The availability correction factor makes the untested assumption that the proportion of dugongs at
the surface is constant across depths, time and activities. Although this is improbable, this
correction factor is likely to be conservative and provides a means of standardising for repeat
surveys of the same area. The availability correction factors for turtles were calculated by
standardising against the number of turtles seen at the surface in a survey of the northern Great
Barrier Reef (Blocks 8Ð13; Marsh and Saalfeld 1989b). The proportion of turtles sighted at the
surface on that survey was the lowest of any survey we have undertaken. The availability
correction factor for turtles is likely to be a considerable underestimate because (i) the correction
factor does not fully account for turtles not visible below the surface, (ii) small turtles are very
difficult to see at the survey altitude, and (iii) turtle sightings are particularly dependent on sea
surface conditions (Marsh and Sinclair 1989a).

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 Availability correction factors were calculated separately from the proportion of dugongs sighted
at the surface in Shark Bay and in NingalooÐExmouth (the proportion of dugongs at the surface did
not vary significantly between Ningaloo and Exmouth: x2 = 0á046, d.f. = 1, P = 0á829). Large
numbers of turtle sightings allowed availability correction factors to be calculated separately for
Shark Bay, Ningaloo and Exmouth. Availability bias was not corrected for dolphins because of the
lack of suitable data to use as a standard.
The surveys were designed primarily to census dugongs. Within that constraint, it was normally
not possible to circle to positively identify small cetaceans, so they have been analysed as a single
group. Similarly, it is difficult to reliably separate green (Chelonia mydas), loggerhead and other
turtles at the survey altitude, so they too have been grouped together.
As the transects were of variable length, the ratio method (Jolly 1969) was used to estimate the
density, population size and associated standard errors for each block. The population estimates
were based on the estimated number of animals of each group of fewer than 10 animals for each
tandem team per transect, calculated with the appropriate corrections for perception and availability
bias and mean group size. The standard errors were adjusted to incorporate the error associated
with each correction factor (Table 2), as outlined by Marsh and Sinclair (1989a). Herds of 10 or
more dugongs are excluded from the calculation of population estimates, and added to the
population estimate as a separate stratum, as suggested by Norton- Griffiths (1978).
The significance of the differences between the abundance of dugongs, dolphins and turtles in
Shark Bay in 1989 and 1994 was tested by ANOVA, both with and without the modal Beaufort sea
state for each transect as the covariate. Blocks and times were treated as fixed factors and
transect as a random factor nested within block. Input data for all analyses were corrected
densities per square kilometre, with each transect contributing one density per survey, based on
the combined corrected counts of both tandem

observer teams. The densities were transformed [log10 (x + 1)] to equalise the error variances. The
differences between surveys of the Exmouth Gulf were examined by two-way ANOVA with transect as the
blocking factor. The differences between surveys of Ningaloo Reef were not compared statistically because
of the differences in the number and lengths of transects.
Sea-surface Temperatures
The best available satellite images of the sea-surface temperatures in the survey areas were obtained for
6 July 1989 (1520 hours, local time) and 23 June 1994 (0717 hours). Temperatures of the sea surface were
derived from AVHRR Bands 4 and 5 by means of an algorithm by McMillin and Crosby (1984) and were
correct to approximately 0á5¡C. Only the top few millimetres of the sea surface is measured, so the recorded
temperatures could be affected by the time of day that the images were acquired.
Results
Shark Bay
Dugongs
During the 1989 survey, 297 dugongs were seen in herds of less than 10. Two large herds
(>40 and >100 dugongs) were also recorded. The mean group size (for herds <10) was 1á48
(± s.e. 0á04). The estimated population was 10146 (± 1665), with an average population density
of 0á71 (± 0.12) dugongs kmÐ2 (Table 3). Calves made up 19% of dugongs sighted.
In the winter of 1994, we counted 290 dugongs on transects, of which 16á6% were calves.
The mean group size was 1á45 (± 0á08; Table 2), and the largest group comprised only seven
dugongs. A ÔherdÕ of at least 200 dugongs was encountered just north-west of Denham, but the
dugongs were dispersed over approximately 10 km 2, so this aggregation was not treated
separately. The estimated population of 10529 (± 1464), and the average density of 0á71
(± 0á10) kmÐ2, were very similar to the 1989 estimates (Table 3).
Comparing the two surveys, there was a significant year-by-block interaction, although
inclusion of Beaufort sea state as a covariate suggests that this effect was marginal (Table 4). In
1989, most dugongs were seen in Blocks 4 and 5 (Fig. 3), and the greatest density (5á1 kmÐ2)
was in Block 5 (Table 3). In 1994, most dugongs, and the greatest density of dugongs
(2á8 kmÐ2), occurred in Block 4 (Fig. 3; Table 3).
In 1989, the distribution of dugongs appeared to be partly determined by water temperature,
with less than 4% of dugongs seen in water colder than 18¡C. This pattern was maintained in
1994, but not as strongly, as 13á8% of dugongs occurred in water colder than 18¡C (Fig. 3).
Turtles
A total of 326 turtles was seen on transects in 1989, compared with 365 in 1994, most of
which could confidently be identified as green turtles. The 1989 population estimate was 6373
(± 710) turtles at an average density of 0á43 km Ð2. In 1994, the population estimate was 8431 (±
758), at an overall density of 0á57 kmÐ2 (Table 3). Comparing surveys, there were significant
effects of year and year-by-block, with or without the inclusion of Beaufort sea state as a
covariate. This suggests that the different distribution of turtles between years is not accountable
to a difference in observation conditions.
In 1989, most turtles were seen in the relatively deep water of Blocks 3, 5 and 6 (eastern
side). In 1994, however, far fewer turtles were seen in these locations. Instead, most were seen
in the shallow waters of the Wooramel Bank and Faure Sill, to the east and south of this area
(Figs 2, 4).
The 1989 distribution of turtles tends to match the pattern in surface water temperature (Fig.
4), with 31á3% of turtles sighted in water colder than 18¡C. In 1994, however, half (50á6%) of
the turtles were seen in water colder than 18¡C. It is unlikely that these differences are due to
major changes in the bayÕs thermal patterns between the time of image capture and aerial survey.
The 1994 image was taken just two days before Blocks 6, 5 and 3 were flown, while in 1989
these blocks were surveyed 4Ð6 days after the image was captured.

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