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1. The whale shark (Rhincodon typus) is a popular focal species within the global marine tourism industry. Although this has contributed to increased protection being granted to the species in several countries, tourism itself can be detrimental to the sharks in the absence of appropriate management. Potential impacts can be mitigated, at least in the short term, by adherence to well-designed interaction guidelines. 2. A burgeoning marine tourism industry based on swimming with whale sharks has developed at Tofo Beach in Mozambique. However, no formal management is currently in place at this site. 3. The behaviour of whale sharks during interactions with boats and swimmers were recorded during 137 commercial snorkelling trips run from Tofo Beach over a 20 month period. Whale sharks were encountered on 87% of trips, which operated year-round. 4. Boat proximity and shark size were significant predictors of avoidance behaviour. No avoidance responses were recorded at 420 m boat distance. 5. The mean in-water interaction time between sharks and swimmers was 8 min 48 s overall. There was a significant decrease in interaction times during encounters where sharks expressed avoidance behaviours, and also in cases where sharks had expressed boat avoidance behaviour before swimmers entered the water. 6. It is suggested that mean encounter times can be extended through adherence to a basic Code of Conduct for operators and swimmers that enforces minimum distances between the sharks, boats and swimmers. Using encounter time as a measure of the 'success' of interactions holds promise, as longer encounters appear to be indicative of lower impacts on sharks while also providing higher customer satisfaction for swimmers.
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Aquatic Conserv: Mar. Freshw. Ecosyst. (2010)
Published online in Wiley Online Library
( DOI: 10.1002/aqc.1149
Developing a Code of Conduct for whale shark interactions
in Mozambique
Manta Ray and Whale Shark Research Centre, Marine Megafauna Association, Praia do Tofo, Inhambane, Mozambique
All Out Africa Research Unit, P.O. Box 153, Lobamba, Swaziland
Eyes on the Horizon, Praia do Tofo, Inhambane, Mozambique
All Out Africa Research Unit, Department of Biological Sciences, University of Swaziland, Private Bag 4, Kwaluseni, Swaziland
1. The whale shark (Rhincodon typus) is a popular focal species within the global marine tourism industry.
Although this has contributed to increased protection being granted to the species in several countries, tourism
itself can be detrimental to the sharks in the absence of appropriate management. Potential impacts can be
mitigated, at least in the short term, by adherence to well-designed interaction guidelines.
2. A burgeoning marine tourism industry based on swimming with whale sharks has developed at Tofo Beach
in Mozambique. However, no formal management is currently in place at this site.
3. The behaviour of whale sharks during interactions with boats and swimmers were recorded during 137
commercial snorkelling trips run from Tofo Beach over a 20 month period. Whale sharks were encountered on
87% of trips, which operated year-round.
4. Boat proximity and shark size were significant predictors of avoidance behaviour. No avoidance responses
were recorded at 420 m boat distance.
5. The mean in-water interaction time between sharks and swimmers was 8 min 48 s overall. There was a
significant decrease in interaction times during encounters where sharks expressed avoidance behaviours, and
also in cases where sharks had expressed boat avoidance behaviour before swimmers entered the water.
6. It is suggested that mean encounter times can be extended through adherence to a basic Code of Conduct for
operators and swimmers that enforces minimum distances between the sharks, boats and swimmers. Using
encounter time as a measure of the ‘success’ of interactions holds promise, as longer encounters appear to be
indicative of lower impacts on sharks while also providing higher customer satisfaction for swimmers.
Copyright r2010 John Wiley & Sons, Ltd.
Received 1 March 2010; Revised 20 August 2010; Accepted 22 August 2010
KEY WORDS: Rhincodon typus; tourism; behavioural ecology; management; conservation
The huge size, placid disposition and planktonic diet of the
whale shark, Rhincodon typus (Smith), has resulted in the
species becoming a popular focus of the global marine tourism
industry (Graham, 2004; Compagno et al., 2005; Catlin and
Jones, 2009). Whale sharks aggregate predictably in certain
tropical and sub-tropical locations to target ephemeral bursts
in local productivity such as plankton blooms (Clark and
Nelson, 1997; Taylor, 2007), fish spawning events (Heyman
et al., 2001) or crab larval release (Meekan et al., 2009). Some
of these areas are, in turn, subject to high tourist visitation
during these periods as people travel to view and swim with the
sharks (Davis et al., 1997; Graham, 2004; Ca
et al., 2007; Quiros, 2007; Rowat and Engelhardt, 2007; Catlin
and Jones, 2009).
Whale shark tourism can give a significant boost to regional
economies, with global revenue estimated to be US$47.5
million in 2004 (Graham, 2004). The annual value of each
of the 106 individual whale sharks identified in Belize was
*Correspondence to: Simon J. Pierce, Manta Ray and Whale Shark Research Centre, Marine Megafauna Association, Praia do Tofo, Inhambane,
Mozambique. E-mail:
Copyright r2010 John Wiley & Sons, Ltd.
estimated to be US$34, 906 in 2002 (Graham, 2004), while
whale shark tour participants spent AU$6.0 million (US$4.5
million) in the Ningaloo region of Western Australia in 2006
(Jones et al., 2009). A high proportion of whale shark tourism
sites are located within developing nations, providing a
compelling economic incentive for these countries to protect
or manage the species (Graham, 2004). Given that whale
sharks have been the subject of large, targeted fisheries in
several countries (Pravin, 2000; Alava et al., 2002; Chen and
Phipps, 2002), the advent of tourism may provide a direct
alternative to extractive harvesting in some areas (Topelko and
Dearden, 2005; Pine et al., 2007).
Shark-related tourism is sometimes viewed as a
controversial activity, particularly when the sharks are
attracted with bait (Topelko and Dearden, 2005; Laroche
et al., 2007). However, whale shark tourism usually involves
either snorkelling or scuba diving with sharks that have not
been actively attracted to observers (Graham and Roberts,
2007; Quiros, 2007) and has thereby largely avoided many of
the perceived negative aspects of the broader industry. In
practice, whale shark tourism has many similarities to in-water
interactions with marine mammals. The latter industry is
comparatively well-studied, and research on a number of large
and small cetacean species has demonstrated that tourism may
have detrimental impacts at an individual, group or population
level (e.g. Bejder and Samuels, 2004; Lusseau, 2005; Bejder
et al., 2006a,b; Stensland and Berggren, 2007). Such impacts
can include both short-term behavioural avoidance and
longer-term population declines (Bejder and Samuels, 2004;
Bejder et al., 2006b). The real or potential impacts of whale
shark tourism are less understood, although certain in-water
swimmer behaviours have been shown to cause short-term
disturbance to the sharks (Quiros, 2007). Increasing tourist
visitation has anecdotally been linked to declines in whale
shark sighting rates in Belize (Graham, 2007) and to other
impacts, such as boat strikes (Ca
´rdenas-Torres et al., 2007;
Rowat et al., 2007), in other locations.
Tofo Beach in southern Mozambique has recently emerged
as an international hot-spot for whale shark encounters.
Although whale sharks have been known to aggregate in the
area for some time (Wolfson, 1986), political circumstances
within the country and a general lack of infrastructure
prevented commercial nature-based marine tourism from
developing until the late 1990s. International tourists now
travel to the region year-round specifically to swim with the
sharks, and at least seven operators currently offer daily
snorkelling trips (‘ocean safaris’). Whale shark tourism has
been identified as a key strategic niche market by the
Mozambican government (Ministe
´rio do Turismo, 2004).
The Mozambican national development plan for tourism
aims to safeguard and manage natural assets to create
desirable tourism products, while simultaneously monitoring
the resources to ensure that tourism itself does not cause
damage (Ministe
´do Turismo, 2004). In most countries where
whale shark tourism is now established, it has proven
necessary to manage interactions through legislation or
voluntary codes of practice to reduce the potential for
negative impacts on the sharks (Davis et al., 1997; Ca
Torres et al., 2007; Quiros, 2007). No legislation pertaining
specifically to whale shark encounters is presently in place in
Mozambique. The objectives of the present study were to
examine the short-term behavioural responses of whale sharks
to boat and swimmer interactions. Using this information,
we provide explicit recommendations for management of this
burgeoning tourism industry through the development of a
best-practice Code of Conduct for interactions with the
Study site and ocean safari procedures
Praia do Tofo (Tofo Beach) (231510S, 351320E) is a small
seaside resort town situated in Inhambane province,
Mozambique, about 400 km north-east of the nation’s
capital, Maputo (Figure 1). At the time of writing,
November 2009, ocean safaris were offered by four operators
at Tofo Beach. Another three operators ran from Barra, which
lies slightly to the north of Tofo, yet searched the same
geographic area for whale sharks. Generally, two or three
operators run ocean safaris on a given day, except during
inclement weather. The main attraction of these ocean safaris
is swimming with whale sharks, although swimmers will also
enter the water opportunistically with bottlenose dolphins
(Tursiops sp.), humpback dolphins (Sousa chinensis) and
manta rays (Manta spp.).
All observations in the present study were made from
a single operator using 8.2 m rigid-hull inflatable craft.
The operational capacity of these vessels was 16–19 people,
usually including three or four staff: a skipper, spotter and a
divemaster/guide. For commercial reasons, trips were rarely
conducted without a minimum of five paying clients. The
majority of trips were made in the middle of the day (between
11 am and 2 pm) to fit in between regular scuba dives and to
facilitate boat-based visual searching for whale sharks through
reduced glare from the sun. Searches were aided on the
majority of trips by the use of a removable spotting chair,
which raised a single observer to approximately 3 m above sea
level to broaden the search corridor. Sharks were sighted by
their dark silhouettes or exposed fins as they swam close to the
Tofo Beach
Tropic of Capricorn
.50 km
Figure 1. Tofo Beach and surrounding coastline. Inset shows the
location of Tofo in Africa.
Copyright r2010 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. (2010)
surface. Search distances were dictated by the number and
duration of animal encounters en-route. Normally an S-shaped
search pattern was driven along the coast from approximately
100 m past the surf-line to 1000 m from shore, with a one-way
travel distance of approximately 6 km. Water depths in this
area range from 5 to 30 m over a predominantly sandy
substrate. Total trip durations were 90–120 min. Trip
characteristics are similar between all the operators working
from Tofo.
Before the trip, a briefing was given onshore to cover
basic safety aspects, boating and whale shark interaction
procedures. A varying amount of environmental interpretation
was provided before and during the trip, but swimmers were
always specifically told not to touch or otherwise harass the
sharks. Upon spotting a shark, the boat was manoeuvred to
place it in or near to the anticipated path of the whale shark.
Swimmers entered the water and snorkelled with the shark
until the encounter was ended by either the shark increasing
speed or diving, swimmers due to fatigue, or operator time
constraints or safety concerns (i.e. when the shark was close to
the surf line).
Data collection and analysis
Observations of whale shark interactions were collected between
January 2008 and August 2009, excluding July and August 2008.
The location of each shark encountered was noted along with
weather conditions and Beaufort sea state. As swimmers entered
the water, the proximity of the shark to the boat, number of
swimmers in the water, estimated underwater visibility and sea
surface temperature (SST) were recorded. The total length of
each shark was estimated, sex was distinguished by the presence
or absence of claspers on the pelvic fins, and the body surface
was examined for wounds or scars.
Behavioural observations for each shark were collected
from both above and within the water. The shark’s initial
response to the boat was categorized as either (a) no reaction;
(b) dive; (c) change of direction; or (d) interaction with the
boat through purposeful close investigation including
occasional ‘bumps’ to the hull or motors. These reactions
were categorized as either avoidance (b) or (c), or no avoidance
(a) or (d). A logistic regression was employed to test whether
the measured variables (in this case SST, sea state, underwater
visibility, sex, size, proximity of boat or scarred/non-scarred)
were significant predictors of behaviour. The effect of boat
proximity on behavioural response was specifically tested using
a Mann–Whitney rank sum test (owing to non-normality of
data), and the influence of the number of boats present
examined using a chi-squared test (w
In the water, a basic ethogram of each shark’s behaviour in
the presence of swimmers was created to record slow
swimming (SS), where the shark remained swimming at a
slow constant rate, or fast swimming (FS) where the shark was
obviously swimming more quickly or accelerated during the
encounter. When the shark was obviously feeding, its
behaviour was categorized as passive feeding (PF), surface
gulping (SG) or underwater gulping (UG). Diving (D) was
noted during the encounter, and behavioural responses to
swimmers such as avoidance (A), change of direction (CD),
banking (B) or interaction (IS), where the shark actively
followed swimmers, were also collated. These behaviours were
summarized as either (1) avoidance, defined as either A, or two
or more of FS, D, CD or B, or (2) no avoidance, comprising all
other behavioural observations. Logistic regression was used
to test whether this binomial response was influenced by
whether the shark was feeding, the number of swimmers in the
water, size and sex of the shark, the presence of external scars,
SST, visibility or sea state.
Total encounter time (in min), defined as the time between
the first swimmer entering the water and the last swimmer
exiting the water, was recorded for each whale shark
interaction. Due to the non-normality of data, the effects of
boat avoidance and in-water avoidance behaviours on
encounter time were specifically assessed using Mann–
Whitney rank sum tests.
In total, 411 interactions with whale sharks were recorded
from 137 trips. Between 0 and 14 shark encounters were
recorded on each trip, with a mean and standard deviation of
3.072.71. At least one shark was sighted on 87.0% of trips.
The number of trips per month where data were collected for
this study varied from 0 to 15. Sharks were sighted in all
months, although there was a significant monthly variation in
the number of sharks sighted per trip (ANOVA on ranks,
17 d.f., Po0.001). The highest mean number of sharks was
recorded in October 2008 (5.572.85 per trip) and the lowest in
August 2009 (0.370.50 per trip) (Figure 2). In both 2008 and
2009, the mean number of sharks observed declined in the
austral winter (Figure 2), indicating that whale sharks move
away from the Tofo region during this season. Sharks were
feeding during 19.4% of encounters.
Behavioural responses to the boat
The proximity of the boat to the shark when swimmers entered
the water ranged from a minimum of o1 m to a maximum of
40 m, with a mean of 7.175.98 m. Where both the proximity of
the shark and its reaction to the boat was recorded (n5264
occasions), an avoidance response was noted during 32.7%
of encounters. Boat proximity (P50.026) and shark size
(P50.015) were the only two significant predictors of
avoidance response among the measured variables (Table 1).
The odds ratio for boat proximity shows that for each extra
metre between the shark and boat, the shark was 0.882 times
less likely to show an avoidance response. Similarly, larger
sharks were less likely to avoid the boat, with each metre of
length decreasing avoidance responses by 0.620 times. Specific
testing of boat proximity effects found mean approach
distances of 5.574.97 m and 7.676.08 m in cases of
avoidance and no avoidance, respectively. No avoidance
responses were noted at proximities 420 m.
Behavioural responses to swimmers
Total encounter times ranged from o1 to 49 min. Excluding
encounters where either the operator ended the encounter
prematurely or swimmers from other boats entered the water
with the shark (n527), the overall mean encounter time was
8 min 48 s78 min 9 s. Swimmers from two or more boats
entered the water with an individual shark on 20 occasions.
The shark exhibited distinct avoidance behaviour upon the
Copyright r2010 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. (2010)
arrival of the second group of swimmers during 60.0% of these
encounters, indicating a significantly higher probability of
avoidance behaviour compared with single-boat encounters
549.84, 1 d.f., P5o0.001).
Behavioural observations were collected for 402 normal
(single boat per shark) in-water interactions. Avoidance
behaviour was noted during 34.6% of these interactions.
Instant diving or abrupt directional changes in response to
swimmer presence (within 30 s of swimmers sighting the shark)
were recorded on 17 occasions (4.8% of in-water interactions),
resulting in encounter times of o1 to 5 min. Encounter time was
negatively related to the expression of avoidance behaviour by
sharks (rank sum test, P50.035), with average encounter times
of 8min 6s78min 2s, and 9min 6s78 min 0 s in cases
where sharks did and did not exhibit avoidance behaviour,
respectively. The only significant predictor of avoidance
behaviour among the measured variables was SST, with each
11C increase in SST corresponding with a 0.842 time decrease in
avoidance responses (Table 1). The number of swimmers per
encounter ranged from 1 to 18, with a median of 11 and a mean
of 10.873.66. There was no obvious relationship between the
number of people in the water and encounter time (linear
regression, 351 d.f., P50.738; Figure 3).
In-water interaction times were also significantly related to
the prior reaction of the shark to the boat. Mean encounter
time was significantly reduced from 8 min 48 s77 min 18 s in
cases where the shark had shown no boat avoidance, to 7 min
0s77 min 39 s when the shark had exhibited an avoidance
reaction (rank sum test, P50.007).
Whale sharks were sighted in every calendar month,
confirming Tofo Beach as having one of the few known
year-round aggregations of the species. There was some
observational evidence for seasonal variation in shark
abundance, with a September/October peak. However, as
trip distance and duration – and therefore the total number of
whale sharks encountered – varied according to the length of
tourist interactions with individual sharks, detailed analysis
of seasonal abundance awaits a more standardized approach.
No. of shark encounters
11 2
Time period
Figure 2. Seasonal variation in shark encounters during 2008 and 2009 (7SD). Numbers above SD lines denote the number of trips undertaken
during that month.
Table 1. Significant predictors of whale shark avoidance behaviours
during interactions with boats and with swimmers, derived from
logistic regression
Boat avoidance Swimmer avoidance
Factor Boat proximity Shark size Sea surface temperature
Pvalue 0.026 0.015 0.006
Odds ratio 0.882 0.620 0.842
Coefficient 0.125 0.478 0.172
Number of swimmers
Encounter time (min)
Figure 3. Mean encounter time (7SD) plotted against the number of
swimmers present.
Copyright r2010 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. (2010)
The overall success rate of tours, i.e. where one or more
sharks were sighted, was 87.0% over the study period. This
figure is higher than the seasonal sighting rates at Ningaloo Reef
in Western Australia (81.6% between 1996 and 2004) (Mau and
Wilson, 2007) and Gladden Spit in Belize (69% between 1998
and 2003) (Graham and Roberts, 2007). The mean number of
whale sharks encountered per trip at Tofo (3.0) was higher than
the mean number of interactions recorded per trip at Ningaloo
between 1996 and 2004, which was 2.6 initially, declining to 1.2
during the period of that study (Mau and Wilson, 2007). This is
a notable point, considering that snorkelling trips at Tofo utilize
boat-based searches along a relatively small length of coast,
whereas in Ningaloo spotter planes are employed to locate the
sharks over a broader area (Mau and Wilson, 2007). The
number of sharks encountered per trip during September and
October at Tofo were similar to in-season rates at Donsol in the
Philippines (6.6 and 8.15 interactions per trip in 2004 and 2005,
respectively) (Quiros, 2007) and Gladden Spit (2 to 6 sharks per
trip between 1998 and 2003) (Graham and Roberts, 2007). The
high trip success rate and mean number of interactions observed
over the period of the present study suggest that Mozambique
has considerable potential as a whale shark tourism destination,
with a ‘product’ that rivals or exceeds more established whale
shark tourism destinations. These results also support aerial
survey data from the South African and southern Mozambican
coasts that had previously recorded relatively high numbers of
whale sharks close to Tofo (Cliff et al., 2007).
Customer satisfaction with in-water interactions with dwarf
minke whales (Balaenoptera acutorostrata) in Australia was
significantly associated with the duration of the encounter
(Valentine et al., 2004), and the same appears to be true
for swimmers with whale sharks (Catlin and Jones, 2009).
In the present study, the expression of short-term avoidance
behaviour by individual whale sharks was linked to a
reduction in encounter time. Therefore, encounter time
appears to have potential as a crude measure of the overall
‘success’ of interactions. For the purposes of the following
discussion, it is explicitly assumed that the primary aim of any
management intervention will be to maximize encounter times,
which is most easily achievable through the minimization of
avoidance behaviour.
A significant link was found between the expression of
avoidance behaviour by sharks and the proximity at which
swimmers entered the water from the vessel. Boat avoidance
behaviour was also associated with shorter encounter times
during ensuing in-water interactions with individual sharks.
This suggests that disturbed sharks either have a heightened
stress response, or in some cases dived before a close interaction
with swimmers could take place. Martin (2007) suggested that
boat avoidance behaviour in whale sharks may be related to
either the low-frequency noise signature of the motors or to a
perceived potential for boat strikes. Scars from small boat
strikes have previously been recorded from Mozambican whale
sharks (Speed et al., 2008), although the observed frequency of
occurrence at Tofo was considerably lower than that reported
from other aggregation sites (Ca
´rdenas-Torres et al., 2007;
Rowat et al., 2007). The presence of scarring was not identified
as a significant predictor of avoidance behaviour in the present
study, although this analysis did not specifically examine
injuries from boats.
Boat avoidance behaviour was not observed at (estimated)
distances 420 m in the present study. These data suggest that
this distance represents a useful initial value for a boat
exclusion radius around sharks. This recommended distance
considerably exceeds the present mean swimmer discharge
distance of slightly over 7 m, suggesting that training
programmes will need to be implemented for skippers and
guides to ensure changes in current behaviour. Given that a
reduction in boat avoidance behaviour is likely to significantly
increase mean in-water encounter times overall, the
application of this new exclusion distance should be
emphasized in training. There is no internationally-applied
boat exclusion radius in use at present, as the situations and
practical realities differ between sites. Code of Conduct
recommendations around the world vary from 5 m in Bahia
de los Angeles, Mexico (Ca
´rdenas-Torres et al., 2007), to 10 m
in Yum Balam, Mexico (Remolina et al., 2007) and 30 m in
Western Australia (DEH, 2005). Furthermore, the high
frequency of avoidance behaviour exhibited by sharks when
interacting with more than one boatload of swimmers clearly
suggests that, as laid out in most national Codes of Conduct,
only one boat should be ‘in contact’ with a shark at one time,
whilst any others maintain a reasonable distance (i.e. outside
the proposed 20 m exclusion radius).
Swimmer interaction times recorded in the present study
represent an intra-site baseline value for future adaptive
management measures. Inter-site interaction times are likely
to be less useful to the formulation of Mozambican
management procedures, as shark behaviour is likely to
change according to feeding strategy and the specific
characteristics of each location. For example, the average
length of interactions in the Philippines, where a higher
proportion of sharks were feeding while observed (in 2005),
was only 3 min (Quiros, 2007). Conversely, mean interaction
times at Ningaloo Reef declined from 27 min in 1996 to 7 min
in 2004, although some interim years remained high, with the
decline possibly influenced by changes in operator procedures
(Mau and Wilson, 2007). Although data on swimmer
behaviours were not collected in the present study, other
studies have shown that maintaining a distance of 3 m from the
body of the shark and 4 m from the tail result in a reduction of
avoidance by sharks. These distances minimize the potential
for accidental touching and also reduce swimmer perceptions
of crowding, thereby improving the quality of the encounter.
Underwater visibility is generally high enough at Tofo to make
these distances practical, unlike in Mexico and the Philippines
where visibility is often poor. In the current study, however,
the physical number of swimmers in the water had no apparent
effect on encounter length, although the mean number of
swimmers was higher than that recommended by most Codes
of Conduct (Quiros, 2007; Remolina et al., 2007; Catlin and
Jones, 2009). It seems likely that, rather than the sheer
presence of swimmers, their behaviour and proximity to the
shark is the important factor to consider in future studies and
management assessments. Sea surface temperature was a
significant predictor of avoidance response, with higher
temperatures associated with decreased encounter times. As
the metabolic rate of ectothermic sharks are strongly affected
by ambient water temperatures (Carlson et al., 2004), this
result may suggest that whale sharks swim faster or are more
responsive to swimmer approach under warmer conditions.
Although the results of the present study show that an
unmanaged tourism industry in Mozambique could have the
potential to cause short-term behavioural modification in
Copyright r2010 John Wiley & Sons, Ltd. Aquatic Conserv: Mar. Freshw. Ecosyst. (2010)
whale sharks, basic mitigation measures should be relatively
simple to implement. The results as shown can reasonably be
taken to approximate the natural behaviour of skippers and
guides, as no formal interaction guidelines were in place during
the study. An increased effort to educate front-line operators
in appropriate interaction techniques is therefore integral to
the success of any new management strategies. Experience
from other countries has shown that instituting accountability
procedures for these staff members is also an important
element, as even relatively low levels of non-compliance can
lead to negative short-term behavioural impacts (Quiros,
2007). Recent studies from Ningaloo Reef in Australia
(Meekan et al., 2006; Holmberg et al., 2008, 2009), Mahe in
the Seychelles (Rowat et al., 2009) and Gladden Spit in Belize
(Graham and Roberts, 2007) have demonstrated that whale
sharks can be temporarily resident or show fidelity to feeding
sites. This suggests the potential for sharks to be repeatedly
exposed to tourist operators, which could result in cumulative
impacts. Quiros (2007) found that sharks sighted for the first
time at Donsol were significantly more likely to exhibit
avoidance behaviour when interacting with swimmers than
sharks encountered repeatedly, which suggests that some
degree of habituation may occur. However, results from
long-term studies on bottlenose dolphins suggest that in some
cases, rather than becoming habituated, sensitive individuals
may simply leave the area (Bejder et al., 2006a,b). This can
lead to long-term population declines even in the absence of
obvious short-term behavioural modification (Bejder et al.,
2006a,b). Given that the length of coast where tours are
conducted in Mozambique is relatively small, a large
proportion of sharks utilizing this area are likely to be
exposed to tourism. This could exacerbate the potential for
negative impacts on Mozambican sharks and highlights the
importance of ongoing monitoring to assess the medium- to
long-term impacts of tourism on whale sharks in this area.
Mozambique plans to attract 4 million tourists annually by
2020 (Ministe
´rio do Turismo, 2004). Such increasing tourist
numbers make it vital to introduce active management for
Mozambique’s whale shark tourism industry to ensure high
quality experiences for swimmers while minimizing detrimental
impacts on the sharks. The Mozambican government is
presently focused on poverty reduction rather than
environmental sustainability. Consequently, realizing the
potential non-consumptive economic value of whale sharks is
likely to be an important management consideration.
However, if this vision of sustainable growth is to be
achieved, iconic tourist species such as whale sharks require
enhanced protection and a dedicated management strategy.
This is the 15th communication of the All Out Africa Research
Unit ( Thanks to all the All Out Africa
volunteers and research coordinators who collected the data
used in the present study, and to Tofo Scuba for providing
support for this research project. SJPs work on this project was
also supported by grants from Project AWARE Foundation
(International) grant 450, and the PADI Foundation grant 23,
and from Casa Barry Lodge. Thanks to Juerg Brunnschweiler
and John Baxter for their helpful comments on the
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... However, the probability of a collision within regions will depend on several factors related to both whale shark behaviors and vessel characteristics. Because shark horizontal swimming speeds were 10 times slower than vessel speeds (SI Appendix, Table S4), whale sharks may be much less able to avoid an oncoming vessel, particularly since if they do exhibit a response it appears slow and limited to until vessels are very close (SI Appendix, Table S5) (25). We were able to determine from tags with depth sensors the proportion of time whale sharks spent near the surface, where they were most at risk of collision (collision zone use, z t ). ...
... We were able to determine from tags with depth sensors the proportion of time whale sharks spent near the surface, where they were most at risk of collision (collision zone use, z t ). However, at the global scale we could not record avoidance responses of sharks to vessels (SI Appendix, Table S5) (25), characterize the type of behavior when at the surface (26), or the range at which oncoming vessels were detected (27,28). Similarly, vessel maneuverability, speed (29), and overall size and draft (24) will also contribute to the risk of collision. ...
... Similarly, vessel maneuverability, speed (29), and overall size and draft (24) will also contribute to the risk of collision. To date there are few studies that explore fine-scale behaviors in sufficient detail to include behavioral aspects into collision-probability estimates for whale sharks (25,30). Given this, our CRI provides an estimate of the risk of collision with large vessels within an area based on the degree of spatiotemporal co-occurrence (hence susceptibility), rather than providing a Table S8). ...
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Significance Global vessel traffic is increasing alongside world economic growth. The potential for rising lethal ship strikes on endangered species of marine megafauna, such as the plankton-feeding whale shark, remains poorly understood since areas of highest overlap are seldom determined across an entire species range. Here we show how satellite tracking whale sharks and large vessel movements globally provides a means to localize high-overlap areas and to determine how collision risk changes in time. Our results point to potential high levels of undetected or unreported ship strikes, which may explain why whale shark populations continue to decline despite protection and low fishing-induced mortality. Collision mitigations in high-collision-risk areas appear necessary to help conserve this iconic species.
... After the discovery of the seasonal presence of the species in Ningaloo Marine Park in the 1980s, where whale shark tourism was first established [12], the whale shark tourism industry has since grown, making substantial contributions to economies as seen in countries like Australia [10], Maldives [11], Mexico [13], Philippines [14,15] and Seychelles [16]. Aside from this, the industry also provides an avenue for scientific research and conservation outcomes [12,[14][15][16][17][18]. ...
... Despite the economic advantages that the industry provides, direct effects of tourism on the host species have been reported by the majority of whale shark behavioural studies within tourist hotspots [14,17,[19][20][21][22][23][24]. Whale sharks react (through banking, diving or changing direction) in the presence of tourists in Ningaloo, Australia [19]. ...
... Similarly, the whale sharks in Donsol, Philippines are likely to be disrupted from feeding, displaying avoidance behaviour during tourist interactions [14]. Whale sharks in Mozambique and Southern Leyte, Philippines, displayed avoidance when tourist vessels were within close proximity [17,24], and during the majority of tourist encounters, suggesting a change in the behaviour of the sharks on at least a short temporal scale [20,24]. These observations, however, are contrary to the behaviours observed in provisioned whale sharks in Oslob, Philippines [21,22]. ...
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The whale shark is the world's largest fish that forms predictable aggregations across its range, many of which support tourism industries. The largest non-captive provisioned whale shark destination globally is at Oslob, Philippines, where more than 500 000 tourists visit yearly. There, the sharks are provisioned daily, year-round, allowing the human–shark interaction in nearshore waters. We used in-water behavioural observations of whale sharks between 2015 and 2017 to understand the relationship between external stimuli and shark behaviour, whether frequency of visits at the site can act as a predictor of behaviour, and the tourist compliance to the code of conduct. Mixed effects models revealed that the number of previous visits at the site was a strong predictor of whale shark behaviour, and that provisioned sharks were less likely to exhibit avoidance. Compliance was poor, with 93% of surveys having people less than 2 m from the animal, highlighting overcrowding of whale sharks at Oslob. Given the behavioural implications to whale sharks highlighted here and the local community's reliance on the tourism industry, it is imperative to improve management strategies to increase tourist compliance and strive for sustainable tourism practices.
... Predictable seasonal aggregations, often associated with high levels of productivity (e.g. spawning events or zooplankton patches containing high densities of shrimp, fish eggs or larvae; Rohner et al., 2015;Tyminski et al., 2015), provide unique opportunities to study these elusive animals (Pierce et al., 2010;Pierce & Norman, 2016;Robinson et al., 2017;Copping et al., 2018). ...
... Because of this, aggregation sites are thought to be key locations for feeding and thermoregulation following deep-water foraging dives (Pierce et al., 2010;Thums et al., 2013;Copping et al., 2018). Strong site fidelity despite disturbance whilst feeding has been recorded at other aggregation sites (Quiros, 2007;Araujo et al., 2017). ...
... Firstly, the differences in behaviour, but not in site use, between injured and non-injured whale sharks suggests an energetic cost to injury, with whale sharks potentially staying at the atoll for extended periods of time to recover from their injuries. As sites thought to be key for thermoregulation and feeding, these aggregation sites may be important locations where recovery and healing can be expedited (Pierce et al., 2010;Thums et al., 2013;Copping et al., 2018). An alternative explanation for the correlation of increasing number and severity of injuries with increasing residency duration could be explained by exposure; residency will probably affect the probability of injury. ...
• Whale sharks collect in predictable seasonal aggregations across the tropics. South Ari Atoll in the Maldives is one of a few aggregation sites where whale sharks can be encountered year‐round. Here, areas with high levels of tourism‐related boating traffic overlap with the whale shark hotspot, increasing the probability of anthropogenic injury. Whale sharks have been reported to remain faithful to this aggregation site following injury, despite the costs of injury and the risk of re‐injury. However, the impacts of injury on site fidelity and residency behaviour are not fully understood. • Encounter data on individual sharks from the Maldives Whale Shark Research Programme database (2006–2018) were analysed to assess the relationship between injury and site fidelity in whale sharks. There was no difference in geographic site use, with injured and non‐injured individuals being encountered in the same areas. However, there were differences in residency timings: injured resident whale sharks (individuals repeatedly encountered over 6 months or longer) spent significantly more time at the atoll and less time absent, and were seen more consistently than non‐injured residents. Increased residency duration, return rate and number of residency periods correlated with increasing injury number. • These differences in behaviour imply a cost to injury, with whale sharks potentially remaining at this site to recover. With boat traffic being concentrated at the aggregation site, injured sharks may be more vulnerable to further injury. Alternatively, these individuals may remain at the atoll despite injury because the benefits gained from this area outweigh the potential costs, with more resident individuals facing greater exposure to anthropogenic threats. These findings highlight the importance of this location and emphasize the need for improved management of anthropogenic activities, particularly boating traffic, at aggregation hotspots to reduce injury rates and any subsequent impacts on behaviour and fitness.
... An increase in evasive behaviours resulting from tourism has also been shown in a variety of animal taxa, including whale sharks, cetaceans, and Galapagos green turtles (Stockin et al., 2008;Pierce et al., 2010;Trave et al., 2017). With increasing tourist pressure, turtles may invest more energy in avoidance behaviours, such as spending less time surfacing and diving, and more time fleeing from approaching tourists (i.e. ...
Anthropogenic activities are increasingly linked to emerging diseases that cause mortality across many taxa. Human interference arising from ecotourism, in particular, can increase the stress levels of wild populations and promote the spread of disease. In Akumal Bay, Mexico, green sea turtles (Chelonia mydas) are increasingly infected with fibropapillomatosis (FP), an infectious disease associated with stress‐induced immunosuppression linked to high human density, which is particularly high in this area because of the increasing number of tourists visiting all year round. To examine whether FP might be associated with behavioural indicators of stress and varying levels of tourist pressure, the behaviour of turtles and the number of tourists were observed through 20‐minute focal sampling periods from May to August 2017. Disease presence and tourist pressure were related to several aspects of turtle behaviour, specifically feeding, resting, vertical movements (i.e. surfacing and diving), and evasive responses. Turtles that had visible FP engaged in fewer feeding periods, vertical movements, and evasive responses. Additionally, with increasing tourist pressure, all turtles spent less time engaging in vertical movements and had more evasive responses. These results suggest that the presence of FP affects green sea turtle behaviour, potentially increasing their exposure to tourists. Sick and healthy turtles appear to react differently to tourists, suggesting that FP changes behavioural responses to tourist pressure. Future management strategies should consider regulating tourist pressure on sick animals to reduce the incidence and progression of FP and other wildlife diseases by limiting access to critical habitats and enforcing code‐of‐conduct compliance from visiting tourists.
... Mozambique (Pierce et al., 2010;Haskell et al., 2015), was introduced among most of the formal tourism operators in 2016 following engagement work by MWSP. These guidelines are currently being added as an inter-ministerial decree into Malagasy Law as an extension of the existing regulation of cetacean tourism. ...
• Between September and December, whale sharks (Rhincodon typus) aggregate in the coastal waters off Nosy Be, an island in north-western Madagascar. Swimming with these sharks has become an important tourism activity, but no formal protection is in place in Madagascar to protect this endangered species from the potential negative effects of tourism or other human impacts. • Boat-based surveys (n = 405) were conducted from tourism vessels from September to December, 2015–2019. For most sightings (98%), whale sharks were sighted while foraging for bait fish at the surface, in association with mackerel tuna (Euthynnus affinis) and seabirds (Sternidae). A total of 408 individual whale sharks were individually photo-identified over this period. All individuals were immature, and 82% of sexed sharks were male. Sharks ranged from 3.0 to 8.0 m in total length (TL), with a mean TL of 5.65 ± 0.94 m (n = 66) for females and 5.46 ± 1.09 m for males (n = 295). • Most sharks (72% of the identified individuals) were only identified once within the study period. Movement modelling showed an open population with a short mean residence time of 7.2 days. Resightings were recorded from up to 12 years apart (2007–2019). Ten sharks were seen in all five seasons during 2015–2019. A basic POPAN mark–recapture model estimated a total population size of 681 (608–763) sharks over the 2015–2019 period. • Nosy Be waters are an important foraging ground for juvenile whale sharks. Sighting data demonstrate that a high proportion of the sharks’ preferred habitat lies outside existing protected areas, but within an identified Key Biodiversity Area. National species-level protection and increased spatial management is warranted to secure the continued presence of whale sharks in this region.
... A voluntary code of conduct was introduced in 2017, based on the Marine Megafauna Foundation's guidelines from Mozambique (from Haskell et al., 2015;Pierce et al., 2010), to codify interactions between whale sharks, boats, and swimmers, and thereby reduce tourism impacts on the whale sharks. The operational code of conduct broadly covers international best-practice standards for whale shark tourism (outlined in ...
Full-text available
Nosy Be in northwestern Madagascar is home to a globally important whale shark (Rhincodon typus) aggregation and a growing whale shark tourism industry. However, whale sharks are not protected in Malagasy waters and are threatened by fisheries bycatch, collisions with vessels, and disturbance from tourism. We used tourist questionnaires (n = 488) to assess the economic value of, and tourist perceptions of, whale shark tourism in Nosy Be from September to December 2019. We also surveyed whale shark tour operators (n = 12) in December 2018 to understand their perceptions of tourism management needs in the region. Results suggest the Nosy Be whale shark tourism industry was worth US$1.5 million for the 3-month 2019 whale shark season. "Dedicated" whale shark divers (i.e., those who came specifically to Nosy Be to swim with whale sharks) spent 55% more money overall and six times the amount individually compared to "casual" whale shark divers. Both tourists and operators supported the protection of whale sharks, with the majority (88.9%) of tourists agreeing that they would choose a tourism destination at which whale sharks are protected. However, tour operators did note significant management issues (e.g., overcrowding, lack of regulations/training), recommending the need to better regulate whale shark tourism and interactions. This study emphasizes the economic rationale for protecting whale sharks in Madagascar to safeguard the emerging marine tourism industry and ensure it is being sustainably managed.
... We also analyzed whether lunges were in the same direction. Furthermore, we recorded whether sharks responded to the vessels present in the area by changing direction, diving, accelerating, inspecting, banking, or vigilance (for definitions see Cubero-Pardo et al. 2011;Montero-Quintana et al. 2020, 2021Pierce et al. 2010;Quiros 2007). Individuals were identified by extracting frames from the videos where the post-gill area of whale sharks is clearly visible and comparing the dot spot pattern using the Interactive Individual Identification System software, I3S (Brooks et al. 2010;Meekan et al. 2006;Speed et al. 2007). ...
Full-text available
Whale sharks are known to feed primarily on zooplankton all over the world; however, recent findings suggest that they also prey on fish using behaviors that have not been fully described. Here, we provide detailed evidence of whale sharks interacting with schools of anchovy on four occasions in Bahia de Los Angeles, Baja California, Mexico. Three of these were predatory interactions in multi-species feeding frenzies, and one was a non-predatory interaction. In predatory interactions, whale sharks exhibited two types of feeding behaviors: (1) stationary suction-feeding, a previously described behavior for whale sharks, and (2) lunge-feeding, which has not been previously described in whale sharks, but has been observed among other large filter feeders, such as rorqual whales. The whale sharks moved simultaneously around the school of anchovy, lunging simultaneously or one after another into the school, with 66% (N = 17) of these lunges occurring in the same direction. In the non-predatory interaction, whale sharks exhibited “sit-and-wait" behavior. The evidence presented here, along with previous observations, suggests that whale sharks may change their feeding strategy from suction to lunge-feeding when other predators corner schooling fish.
... Similar to our results, the mean counts of Galapagos (Carcharhinus galapagensis), sandbar (Carcharhinus plumbeus), and tiger sharks at a dive site in Hawaii did not change when one or two boats were present (Meyer et al., 2009). However, whale sharks (Rhincodon typus) tended to show a higher frequency of avoidance responses when more boats were present (Pierce et al., 2010). Such patterns were not found in the response of great hammerhead sharks in Bimini, where philopatric sharks were not influenced by the number of boats and new sharks were even more likely to be present when boat numbers increased (see below). ...
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Wildlife provisioning is popular, economically valuable, and a rapidly growing part of marine tourism, with great potential to benefit conservation. However, it remains controversial due to limited understanding of its implications on the behavior and ecology of target species. In this study, we modeled how various abiotic and biotic factors influenced great hammerhead sharks' (Sphyrna mokarran) use of a recreational dive site in Bimini, the Bahamas, where shark-feeding has been conducted since 2012. Further, we calculated bioenergetic models to estimate their daily ration and examined if individual sharks fulfilled their daily energetic requirements from food uptake during dives. Between December 2016 and May 2017, we collected data during 104 provisioning dives in collaboration with a local dive operator. Twenty-eight individual great hammerhead sharks were observed, 11 were philopatric (i.e., identified at the dive site in previous years), and 17 were new (i.e., identified at the dive site for the first time during this study) individuals. On average, four sharks were observed daily, occasionally up to nine individuals, with some individuals spending more than 2 h attending each dive, consuming up to 4.75 kg of provisioned food per dive and returning repeatedly throughout the study period. When we grouped sharks based on their previous experience of the dive site (i.e., philopatric vs. new sharks), we found significantly higher attendance indices, i.e., the number of attended dives divided by the total number dives, and longer presence times during dives in philopatric sharks and different responses toward the number of boats and conspecifics between the two groups. Overall, great hammerhead sharks increased their bait uptake during longer dives and when more boats were present at the dive site. Finally, nine of 12 provisioned great hammerhead sharks were regularly able to fuel their daily energetic requirements from provisioned food alone, with two sharks doing so on 77.8% of all dives. Our study provides insights into how large-bodied marine predators react toward wildlife tourism associated provisioning and allows further discussion about daily energy uptake during provisioning dives, its potential impacts on the ecological role of the target species and associated management measures.
... Many countries where whale sharks are found conduct such tourism activities to create income for the local people. However, this tourism must be well managed otherwise it can have negative impacts including changes in whale shark behavior (Pierce et al. 2010;Fitzpatrick et at. 2011;Haskell et al. 2015). ...
Djunaidi A, Jompa J, Nadiarti N, Bahar A, Tilahunga SD, Lilienfeld D, Hani MS. 2020. Analysis of two whale shark watching destinations in Indonesia: status and ecotourism potential. Biodiversitas 21: 4511-4523. Botubarani in Gorontalo, on the northern coast of Tomini Bay and Labuhan Jambu in Teluk Saleh, Sumbawa recently became whale shark tourism destinations in Indonesia. Both sites offer visitors opportunities to interact with whale sharks, either watching from canoes or snorkelling and diving. In this study, we investigated the status and ecotourism potential of whale shark watching as a form of marine wildlife tourism development in eastern Indonesia by examining the existing operations in both sites. Data were collected using questionnaires administered to local community members and visitors to understand their perceptions, and how this opportunity was valued. Secondary data were collected to explore and understand influencing factors. Findings revealed that whale sharks commonly emerge near bagan lift-nets and other interactive areas as a response to the presence of their favorite prey, anchovies, and rebon shrimp. In general, whale sharks most commonly visit bagan lift nets and can be found in the interaction areas in the morning and stay for couple of hours before swimming away to deeper waters. Out of 119 community respondents in Labuhan Jambu village (Teluk Saleh) and Botubarani, Gorontalo, 80.7% knew about the occurrence of whale sharks in their locality. Of the 111 visitor respondents, 67.6% knew about the whale shark watching sites from word of mouth. The study results also suggest a correlation between chlorophyll-a concentration, anchovies, rebon shrimp and whale sharks. Regarding management approaches to this type of tourism, a local community-based management model is strongly recommended, with 87.4% of respondents showing interest in this approach. The roles and responsibility of local government in developing such tourism is very significant, especially with respect to facilitating appropriate competency within the local community throughout training and education, as suggested by nearly 26% of respondents.
The aim of this study is to provide preliminary observations on the microanatomy of Rhincodon typus skin by means of histology and electron microscopy analyses. Skin biopsies were obtained from a deceased juvenile male shark (548 cm total length) stranded in La Paz, Mexico during February 2018. Our results evidenced the basic structure of the dermal denticles in the epidermis of the trunk of the shark, as well as the composition of the connective tissue in the hypodermis. Histological images of the hypodermis showed a high concentration of collagen fibres, formed by a large number of fine and wavy fibres of compact shape and little intercellular substance. This article is protected by copyright. All rights reserved.
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Occurrence of whale sharks in India was considered a rarity until recently. Details of whale sharks that have landed so far in the Indian coasts are summarized. The whale shark has become a regular fishery in successive years off Gujarat coast for its meat, fins, liver, skin and cartilage. Over 1000 whale sharks have been hunted off Saurashtra coast during 1998. The present paper gives an account on the year-wise, season-wise, state-wise, depth-wise and gearwise distribution and occurrence of whale sharks, based on incidental landings and capture, in Indian coastal waters from 1889 to 1998. The threat to the whale shark due to commercial exploitation, its utilization and the need for conservation is discussed.
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Capture-mark-recapture (CMR) data from Ningaloo Marine Park (NMP) in Western Australia have recently been used to study the population dynamics of the local whale shark aggregation. Because nascent research efforts at other aggregation points look to NMP as a model, further analysis of existing modeling approaches is important. We have expanded upon previous studies of NMP whale sharks by estimating CMR survival and recruitment rates as functions of average total length (TL). Our analysis suggests a decline in reported values of TL coincident with marginally increasing abundance among sharks sighted in more than one year ('returning') from 1995 to 2008. We found a positive, average returning recruitment rate (χ) of 1.07 yr-1 (0.99 to 1.15, 95% CI); smaller individuals contributed in larger numbers to recruitment, allowing for population growth accompanied by a decline in median size. We subseguently explored intraseasonal population dynamics with the Open Robust Design (ORD) model structure. Our best-fit model estimated modestly increasing annual abundances between 107 (95 % CI = 90 to 124) and 159 (95% CI = 127 to 190) for 2004 to 2007, suggesting a short-term increase in total annual abundance. The ORD also estimated an average residency time of 33 d (95% CI = 31 to 39) and biweekly entry profiles into the study area. Overall, our techniques demonstrate how large aggregations of the species can be modeled to better understand short- and long-term population trends. These results also show the direct scientific benefit from the development of an online, collaborative data management system to increase collection of sighting data for a rare species in conjunction with ecotourism activity.
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Ecotourism operations which provide food to large predators have the potential to negatively affect their target species, by conditioning them to associate humans with food, or by generally altering their behavioural patterns, This latter effect could have potentially detrimental consequences for the ecosystem inhabited by the predator, because any behavioural changes could affect the species with which they interact. We present the results of an experimental study conducted from June to October 2004, which examined the effects of provisioning ecotourism on the behaviour of white sharks around a seal colony on a small island in South Africa. Although ecotourism activity had an effect on the behaviour of some sharks, this was relatively minor, and the majority of sharks showed little interest in the food rewards on offer. It is unlikely that conditioning would occur from the amount of ecotourism activity tested, because even those sharks identified supplying most of the data presented here (which may be more strongly predisposed towards conditioning, as their persistence around the boat is what allowed them to be identified) showed. a nearly ubiquitous trend of decreasing response with time. Furthermore, even the sharks frequently acquiring food rewards typically stopped responding after several interactions. Consequently, moderate levels of ecotourism probably have only a minor impact on the behaviour of white sharks, and are therefore unlikely to create behavioural effects at the ecosystem level.
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A population of bottlenose dolphins inhabits 7 of the 14 fjords that compose Fiordland, New Zealand. One of these fjords, Milford Sound, supports a large tourism industry that results in intense boat traffic. Bottlenose dolphins regularly visited Milford Sound and tour boats interacted with them during these visits. I studied the factors affecting the frequency of visits to Milford Sound by relating the residency pattern of dolphins in this fjord to oceanographic parameters and variations in boat traffic between December 1999 and February 2002. Boat traffic was the only variable that could explain the frequency of dolphin visits to Milford Sound. Dolphins spent less time in Milford Sound during seasons of intense boat traffic. Moreover, when dolphins visited this fjord, they spent more time at the entrance of the fjord when boat traffic was intense, out of the reach of tour boats. It seems that dolphins avoid Milford Sound when traffic is heavy. This avoidance could have long-term implications for the demography of the population.
Introduction Despite the ecological significance of elasmobranchs as top-level predators in most marine ecosystems (Cortés, 1999), information on their energetics and metabolism is meager. Metabolism is an important component of an organism’s daily energy budget and may account for its greatest, yet most variable proportion (Lowe, 2001). It was hypothesized that sharks had lower metabolic rates than comparable teleosts because most of the original work on the metabolic rate of sharks focused on relatively inactive, cooler-water sharks such as spotted dogfish, Scyliorhinus canicula (Piiper and Schumann, 1967; Metcalf and Butler, 1984) and spiny dogfish, Squalus acanthias (Brett and Blackburn, 1978). Over time, better techniques have evolved that allow study of more active elasmobranch species that were typically considered difficult to work with in captivity. These advances in technology have expanded our knowledge of ecology, activity level, morphology, cellular physiology, and kinematics of elasmobranchs that exhibit a wide range of lifestyles, indicating that elasmobranchs have metabolic rates comparable to teleost fishes of similar size and lifestyle.
We investigated the behavioural changes of Indo-Pacific bottlenose dolphins Tursiops aduncus in response to boat-based tourism at both group and individual levels. The behaviour, movement and dive patterns of nursing females off the south coast of Zanzibar were investigated between January and March 2000 to 2002 and statistical comparisons were made between observations made at different levels of tourist activity. Behavioural data was collected during boat surveys using scan sampling of groups and focal individual follows of 5 female dolphins with calves. The movement patterns of dolphin groups were not affected by the presence of a few (1 to 2) tourist boats without swimmers. However, the groups displayed a significantly larger proportion of erratic (non-directional) movements as tourist activities increased and when swimmers were present. The proportion of active, peduncle, tail-out and porpoise dives also increased as tourist activity increased. Further, females travelled more frequently as tourist activities increased; this may have a negative effect on the time available for females to nurse their calves. Intense non-regulated dolphin tourism in this area may lead to a shift in habitat use by nursing females, and the apparent changes in dolphin behaviour due to the increased levels of tourism may ultimately reduce fitness at both individual and population levels. We urge that the guidelines already issued by the Department of Fisheries and Marine Products, Zanzibar, be implemented and complied with as a first important step towards sustainable dolphin tourism.