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Monitoring the effects of tourism on whale shark Rhincodon typus behaviour in Mozambique

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The whale shark Rhincodon typus is a popular focal species in the marine tourism industry. We analysed 689 encounters with at least 142 individual sharks during 2008–2010 to assess their behaviour in the presence of swimmers at Tofo Beach, Mozambique. Sharks varied in size (estimated 3.0–9.5 m total length) and the majority (74%) were males. The sharks displayed avoidance behaviours during 64.7% of encounters. Encounter duration decreased significantly, from 12 minutes 37 s with undisturbed sharks to 8 minutes 25 s when sharks expressed avoidance behaviours, indicating that interactions with tourists affected the sharks’ short-term behaviour. However, during the 2.5-year study period we found no trend in the mean encounter duration, the overall expression of avoidance behaviour or the likelihood of an individual shark exhibiting avoidance behaviours. Potential effects of tourism may be mitigated by the non-breeding status and transient behaviour of sharks at this aggregation site.
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Monitoring the effects of tourism on whale shark
Rhincodon typus behaviour in Mozambique
PETER J. HASKELL,ANDREW MCGOWAN,ANNA WESTLING,ADRIANA MÉNDEZ-
JIMÉNEZ,CHRISTOPH A. ROHNER,KYM COLLINS,MARCELA ROSERO-CAICEDO
JODI SALMOND,ARA MONADJEM,ANDREA D. MARSHALL and S IMON J. PIERCE
Abstract The whale shark Rhincodon typus is a popular
focal species in the marine tourism industry. We analysed
 encounters with at least  individual sharks during
 to assess their behaviour in the presence of
swimmers at Tofo Beach, Mozambique. Sharks varied in
size (estimated ..m total length) and the majority
(%) were males. The sharks displayed avoidance behav-
iours during .% of encounters. Encounter duration de-
creased significantly, from  minutes  s with
undisturbed sharks to minutes  s when sharks expressed
avoidance behaviours, indicating that interactions with
tourists affected the sharksshort-term behaviour.
However, during the .-year study period we found no
trend in the mean encounter duration, the overall expres-
sion of avoidance behaviour or the likelihood of an individ-
ual shark exhibiting avoidance behaviours. Potential effects
of tourism may be mitigated by the non-breeding status and
transient behaviour of sharks at this aggregation site.
Keywords Behavioural observations, ecotourism, Mozambique,
Rhincodon typus, risk assessment, tourism management, whale
shark
This paper contains supplementary material that can be found
online at http://journals.cambridge.org
Introduction
Viewing sharks in their natural setting is a popular tour-
ism activity (Gallagher & Hammerschlag, ) and
the income accrued creates an incentive to manage these
charismatic species as a non-consumptive resource
(Brunnschweiler, ; Clua et al., ; Vianna et al.,
). However, studies on elasmobranchs have documen-
ted situations in which marine tourism has negative effects,
including behavioural changes and increased energetic costs
(Pierce et al., ; Fitzpatrick et al., ). Improved under-
standing of the actual or potential effects of tourism inter-
actions is important for mitigating or avoiding longer-term
effects and ultimately safeguarding employment and the
tourism infrastructure.
The whale shark Rhincodon typus is the largest fish and
an iconic species for tourism (Gallagher & Hammerschlag,
). Commercial whale shark interaction tours began in
 in Western Australia after a seasonal aggregation of
the species was discovered within the Ningaloo Marine
Park (Davis et al., ). In  the whale shark tourism
industry at this site was valued at AUD million annually
(Catlin et al., ). Whale shark tourism industries have
now developed at several locations in all three tropical
oceans and are mostly based on the presence of predictable
seasonal feeding aggregations of sharks exploiting ephem-
eral bursts in local productivity, such as mass fish or coral
spawning events (Taylor, ; de la Parra Venegas et al.,
). The global revenue from whale shark tourism was
provisionally estimated to be USD  million in 
(Graham, ).
Whale sharks are categorized as Vulnerable on the IUCN
Red List (Norman, ). Although the economic value or
potential of whale shark tourism has helped to justify legal
protection for the species in some countries, concerns have
also been raised that specialist tourism industries could
negatively affect the sharks. Sightings of whale sharks at
Gladden Spit in Belize declined during 
(Graham & Roberts, ) and, based on anecdotal reports
from guides, remained low at least until  (Graham,
). Graham () suggested that the rapid increase in
diver numbers at this site may have led to disturbance of
snapper spawning behaviour (the main driver of whale
shark presence), and the whale sharks themselves, although
a dedicated study on disturbance by divers did not identify a
PETER J. HASKELL* and ANDREW MCGOWAN School of Biosciences, University of
Exeter, Penryn, UK
ANNA WESTLING,ADRIANA MÉNDEZ-JIMÉNEZ,KYM COLLINS,MARCELA ROSERO-
CAICEDO and JODI SALMOND All Out Africa Research Unit, Lobamba, Swaziland
CHRISTOPH A. ROHNER,ANDREA D. MARSHALLand SIMON J. PIERCE§
(Corresponding author) Marine Megafauna Foundation, Tofo Beach,
Inhambane, Mozambique. E-mail simon@marinemegafauna.org
ARA MONADJEM All Out Africa Research Unit, Department of Biological Sciences,
University of Swaziland, Kwaluseni, Swaziland
*Also at: Marine Megafauna Foundation, Tofo Beach, Inhambane, Mozambique
Also at: School of Geography, Planning and Environmental Management, The
University of Queensland, St Lucia, Australia, and Climate Adaptation Flagship,
CSIRO Marine and Atmospheric Research, EcoScience Precinct, Dutton Park,
Australia
Also at: Wild Me, Tofo Beach, Inhambane, Mozambique
§Also at: All Out Africa Research Unit, Lobamba, Swaziland
Received February . Revision requested  April .
Accepted September .
©2014 Fauna & Flora International,
Oryx
, Page 1 of 8 doi:10.1017/S0030605313001257
direct effect on the sharks (Heyman et al., ). Propeller
injuries from small boats have also been observed on whale
sharks at several aggregation sites (Rowat et al., ; Speed
et al., ). Studies of the short-term behavioural responses
of sharks to tourists and boats at Ningaloo Reef (Norman,
), Donsol in the Philippines (Quiros, ) and Tofo
Beach in Mozambique (Pierce et al., ) have revealed
that sharks routinely display avoidance behaviours, includ-
ing banking, eye-rolling, fast swimming and diving, in
response to close approaches by swimmers or boats.
The ecological, social and economic sustainability of
whale shark tourism at Ningaloo Reef was reviewed by
Mau (). Although long-term empirical data on whale
shark behaviour in the area were not available, the industry
was judged to be ecologically sustainable. This was based on
the lack of observed interruption of feeding behaviour,
which may take place largely at night or at least outside tour-
ist interaction times (Taylor, ), the regular re-sightings
of philopatric sharks (Holmberg et al., ,), and the
lack of reproductive behaviour observed in this juvenile
male-biased population (Meekan et al., ; Norman &
Stevens, ). However, concerns were raised about the
potential for injury from boat strikes (Speed et al., )
and an apparent decline in mean size over time
(Bradshaw et al., ). Aside from Ningaloo Reef, where
the sharks have been relatively well studied, the longer-term
sustainability of whale shark tourism has not been consid-
ered explicitly.
Here we examine the tourism industry at Tofo Beach
(Praia do Tofo) in Mozambique, an international hotspot
for whale shark encounters (Pierce et al., ). Whale
shark interactions are a key attraction for international
divers visiting the country but a lack of official management
has raised questions about the sustainability of the industry
(Pierce et al., ; Tibiriçá et al., ), particularly after a
significant decline in sightings during  (Rohner
et al., ). Previous work at Tofo Beach has examined
how interactions can be managed to minimize the potential
for short-term negative effects on the sharks (Pierce et al.,
). Here we extend that dataset to evaluate whether lon-
gitudinal encounter data reveal evidence of an increasing
frequency of avoidance behaviours overall and whether
individual-based analyses show evidence of changing avoid-
ance behaviours over time. We consider the implications of
these data and evaluate the longer-term implications for the
ecology of whale sharks at this site.
Methods
The village of Tofo Beach is situated in the Inhambane prov-
ince of Mozambique, c.  km north-east of the capital
city, Maputo (Fig. ). A full site overview and description
of the commercial whale shark tourism industry in Tofo
are provided by Pierce et al. (). Tourism operators
offer daily -hour snorkelling trips that aim to locate
whale sharks and other marine megafauna. Vessels typically
survey a km stretch of coastline south of Tofo, between the
surf line and c. , m from the shore, in waters c.  m
deep. Sharks are located through visual inspection of the
water surface, where it is often possible to spot their dark
silhouettes or exposed fins. The boat is then positioned in
relation to the sharks direction of travel and clients enter
the water to interact with the shark.
We collected data during January June ,
except during July and August  when no sampling
trips took place. Upon locating a whale shark, observers en-
tered the water alongside clients and recorded the total
number of swimmers and environmental characteristics,
including weather conditions (categorized as sunny, slightly
overcast, overcast or raining), Beaufort sea state and under-
water visibility. The total length of the shark was estimated
visually (Rohner et al., ) and the sex identified by the
presence or absence of claspers on the pelvic fins. The
presence and location of any injuries or scars were noted
and categorized post hoc as either major or minor (Speed
et al., ). A basic ethogram of each sharks behaviour
in the presence of swimmers was produced to record slow
swimming, equating to normal behaviour; fast swimming,
where there was an obvious increase in the sharks tail-beat
frequency; diving, where the shark dived away from the sur-
face; banking, where the shark rolled its back towards swim-
mers; changing direction, where the shark altered its
direction of swimming in the presence of swimmers; and
any other obvious avoidance behaviours, such as the violent
shudder reported by Quiros (). Observations of feeding
behaviour were also noted when they occurred. The total
encounter duration, defined as the time between the first
swimmer entering the water and the last swimmer returning
to the boat, was recorded (in minutes) following each
interaction. When possible, standardized identification
FIG. 1 Tofo Beach, Mozambique. The rectangle on the inset
shows the location of the main map in south-east Africa.
2 P. J. Haskell et al.
©2014 Fauna & Flora International,
Oryx
,18
photographs of both the left and right flanks of each shark
were taken and uploaded to the global Wildbook for Whale
Sharks photo-identification library (Arzoumanian et al.,
; Marshall & Pierce, ). Each of these encounters
was then assigned to a new or previously identified shark
in the library.
It was difficult to establish whether behaviours exhibited
by whale sharks in the presence of swimmers would also
occur in a natural, undisturbed setting. Within-effect com-
parisons (i.e. behaviour in the presence of swimmers) were
therefore employed instead of attempting to determine
cause-and-effect relationships. This approach is useful in
cases where there are no baseline or control data, and
focuses on assessing behavioural responses under
gradations of the effect (Bejder & Samuels, ). Fast
swimming, diving, banking and changing direction are typ-
ically classified as avoidance behaviours in whale sharks
(Norman, ; Quiros, ; Pierce et al., ). When
these or other obvious avoidance behaviours were observed
during an interaction they were each assigned a score of one.
The total score at the end of each encounter was thus equal
to the total number of avoidance behaviours observed. All
statistical analyses were carried out using Rv...(R
Development Core Team, ), with MASS and nlme.
The relationship between avoidance score and encounter
duration was tested using a generalized linear mixed model
approach. Encounter duration, log transformed to ensure
that error structure was normally distributed, was used as
the response variable. Avoidance score was used as the
explanatory variable and shark identification was added as
a random effect to avoid pseudo replication. Given the sig-
nificant relationship between avoidance score and encoun-
ter duration (see results; Pierce et al., ), encounter
duration was used in all further analyses as a proxy for
avoidance. A further model was fitted to determine which
variables significantly affected encounter duration.
Encounter duration was entered as the response variable
and tested against nine main effect explanatory variables:
whale shark variables (size, sex, presence/absence of scar-
ring or feeding behaviour), environmental variables
(weather, sea state () and underwater visibility), number
of swimmers present during an encounter, and the Julian
day (Day =January ). We tested second-order inter-
actions between all combinations of whale shark variables;
all combinations of environmental variables; size, sex and
scarring and all environmental variables; and between the
number of swimmers and underwater visibility. Year and
month of encounter were entered as nested random effects.
Only encounters in which shark identity had been con-
firmed were used in the analysis; a single interaction was
randomly selected from those individuals for which mul-
tiple encounters had been recorded. We used a generalized
linear model to test for a significant relationship between the
amount of scarring on an individual and its total length.
A generalized linear model was fitted, using the number
of encounters as the response variable and search effort
(in hours) and year as explanatory variables, to establish
whether encounter rate changed over the course of the
study. As the dataset ended in June , a second model
was fitted to -month periods, using the number of encoun-
ters as the response variable and search effort and -month
block as explanatory variables. A quasi-poisson error struc-
ture was used for both analyses because of overdispersion of
residual deviance. A third model was used to test for a
change in encounter duration during the study period,
using log-transformed encounter duration as the response
variable and month and year as explanatory variables. We
used χ
to test for a population-level increase in avoidance
response over the five -month periods. A generalized linear
mixed effects model was used to test whether the likelihood
of avoidance was related to the number of previous encoun-
ters in which the whale shark had been successfully identi-
fied and its behaviours recorded over the study period.
We used the Wildbook library to establish how many previ-
ous encounters had been recorded for each individual shark.
A binomial score for avoidance was used as the response
variable, encounter number as the explanatory variable,
and shark identity as a random factor. Significance was
accepted at the % confidence interval in all analyses.
Results
We recorded a total of  whale shark encounters from 
trips during January June , with a total search
effort of .hours. The mean encounter rate was . ±SD
. per trip, with  encounters recorded on each trip,
equating to . ±SD . sharks per hour of search effort.
At least one shark was sighted on .% of trips. There
was no significant trend in sightings over the course of the
study (linear regression, df = ,P=.;Fig. ). A total of
 sharks were positively identified using the Wildbook
photo-identification library. Of  sexed sharks,  were
female (.%) and  were male (.%), which was
FIG. 2 Mean (±SD) number of whale sharks Rhincodon typus
sighted per daily trip during the -month sampling period.
Numbers above the bars denote the number of sampling days in
that month.
Tourism and whale sharks 3
©2014 Fauna & Flora International,
Oryx
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significantly different from a :sex ratio (χ
=.,
P#.). The mean total length was . ±SD . m
(range .m) for females and . ±SD . m (range
.m) for males (Fig. ), with no significant difference
between the sexes (t-test, t=.,df=,P=.).
The total number of encounters with identified sharks
(counted as the number of days on which interactions
with a particular shark were recorded) was over the
course of the study, although the inclusion of additional
data from the Wildbook library showed that % of the indi-
vidual sharks considered in this study had been exposed to
swimmers prior to the start of this work in . Overall,
sharks were encountered on a mean of . ±SD . occa-
sions, with up to  encounters per individual (Fig. ). The
reliance on suitable photographs having been submitted to
the library means that these figures represent the minimum
number of previous encounters. Scarring was observed on
.% of identified individuals and whale sharks were ob-
served feeding during .% of all encounters.
The mean duration of encounter was minutes  s±
SD minutes  s(n=). One or more avoidance re-
sponses were recorded during .% of encounters.
Evaluation of  encounters with  identified sharks in-
dicated that encounter duration was significantly related to
expressed avoidance response (generalized linear mixed
model, χ
=.,P=.). The mean duration of
encounter when a shark showed no avoidance was  min-
utes  s±SD minutes s(n=), which is significantly
longer than encounters with sharks that exhibited avoid-
ance (generalized linear mixed model, χ
=.,
P,.). There was no significant difference in the
duration of encounters where one or more avoidance
behaviours were expressed (generalized linear mixed
model, χ
=.,P=.); mean duration of such en-
counters was minutes  s±SD minutes  s(n=).
A binomial response (avoidance or no avoidance) was
therefore used in subsequent analyses.
All potential explanatory variables were present in 
encounters and, after controlling for multiple encounters
with individual sharks,  encounters were available to
test for other significant variables affecting encounter dur-
ation. A linear mixed effects model, with year and month
retained as nested random effects, demonstrated that scar-
ring was highly significant (linear mixed effects, χ
=.,
P=.). Encounters with individuals that had no appar-
ent scarring were significantly shorter than with individuals
that had some degree of scarring (linear mixed effects,
χ
=.,P=.); the severity of the scarring did not sig-
nificantly affect encounter duration (linear mixed effects,
χ
=.,P=.). There was no significant relationship
between the total length of an individual and the degree of
scarring observed (ANOVA, F
(,)
=.,P=.).
Encounters were significantly longer when sharks were
feeding (linear mixed effects, χ
=.,P=.) and
when more swimmers were present (linear mixed effects,
χ
=.,P=.).
The number of whale sharks encountered was not
correlated with search effort between years (ANOVA,
F
(,)
=.,P=.). There was a significant difference
in encounter rates between years (ANOVA, F
(,)
=.,
P=.) but analysis of -month blocks indicates that the
source of this difference was the second half of , for
which July and August were not included (ANOVA,
F
(,)
=.,P#.). Encounter rate was generally
low during the austral winter, so excluding these months
may have affected this result. No other variables were
significant. There was no significant difference in the
frequency of avoidance behaviours expressed between the
five -month periods (χ
=.,P=.). Encounter
duration was significantly shorter in  (minutes s±
SD minutes  s) than in  ( minutes  s±SD
minutes s) and  ( minutes  s±SD minutes
 s; ANOVA, F
(,)
=.,P#.). A total of 
encounters with  whale sharks were analysed to establish
that the number of previous encounters with swimmers did
not affect the likelihood that a whale shark would display
FIG. 3 Frequency distribution of the estimated lengths of
photo-identified sharks.
FIG. 4 The total number of sightings (unique encounter days) for
each photo-identified shark.
4 P. J. Haskell et al.
©2014 Fauna & Flora International,
Oryx
,18
avoidance behaviour (generalized linear mixed model,
χ
=.,P=.).
Discussion
We found no significant longitudinal effects on encounter
duration or the raw proportion of avoidance behaviours ex-
pressed over the -month study period. Furthermore, there
was no significant relationship between the number of
previous tourist encounters and the likelihood of avoidance
behaviours being displayed by individual sharks. At this
stage there is no evidence of short-term avoidance beha-
viours translating into medium- to longer-term behavioural
change among the sharks present in Mozambican waters.
The potential negative effects of tourism may be ameli-
orated by the ecology and population structure of whale
sharks in Mozambique. The majority of sharks encountered
were male (%). Maturity in male sharks occurs when they
reach a total length of c. m, based on records from
Ningaloo Reef (Norman & Stevens, ), and in female
sharks at a total length of ..m, based on South
African and other records (Beckley et al., ; Norman
& Stevens, ). Given that no sharks of total length
..mor,m were encountered, most sharks were
immature and were probably several years old at the time
of first sighting (Wintner, ). Therefore, tourism at
this site is unlikely to interfere with reproduction, if it
does in fact occur in these waters. The majority (%) of
sharks had only one or two encounters recorded on the
Wildbook photo-identification library by  June 
(with the earliest record from ), including from this
study, suggesting that many of the sharks are transient to
the area, as noted in other aggregations (Holmberg et al.,
; Rowat et al., ; Fox et al., ). There was no evi-
dence of a learnt avoidance behaviour amongst individual
sharks and, based on data from the Philippines (Quiros,
), a degree of habituation may be more likely to occur.
The percentage of whale sharks observed feeding during
daylight hours at Tofo (.%) is comparable to the rates re-
ported at Donsol in the Philippines (%in and %in
; Quiros, ) but substantially lower than the % re-
ported from Bahía de Los Angeles, Mexico, where research-
ers targeted feeding whale sharks (Nelson & Eckert, ).
Contrary to results from the Philippines, where feeding
sharks were . times more likely to exhibit a dive response
in the presence of swimmers (Quiros, ), encounter
durations in this study were significantly longer when feed-
ing behaviour was observed. Whale sharks are flexible in
their foraging strategies, depending on prey type and verti-
cal distribution (Graham et al., ; Nelson & Eckert, ;
Taylor, ; Brunnschweiler et al., ; Motta et al., ),
so the effects of tourism on feeding are likely to be
context-specific.
Whale shark encounters were spatially aggregated along
km of coast immediately south of Tofo Beach. Although
the sharks are also sighted more widely along the
Mozambican coast (Cliff et al., ), the small size of the
primary aggregation area and its almost complete daily
search coverage by commercial operators suggest the possi-
bility of displacement if whale sharks have negative inter-
actions with swimmers or boats. Over half (%) of
identified sharks had some form of scarring, although not
all of these were from anthropogenic sources such as propel-
ler strikes. This percentage is lower than that recorded in
Djibouti, where propeller or boat strike scars were observed
on % of identified sharks (Rowat et al., ), but empha-
sizes the importance of working with skippers to ensure that
a safe minimum distance from the sharks is maintained
(Pierce et al., ). Although tourism operators have not
perceived a shift in whale shark distribution, the lack of
standardized spatial sampling coverage precludes a defini-
tive assessment.
Our results support the use of encounter duration as a
means of measuring the effect of swimmer presence on
whale shark behaviour, as sharks that are disturbed will rou-
tinely end an encounter by diving or increasing their swim-
ming speed. However, it is important to note that encounter
duration, although a useful proxy of avoidance, may only be
able to provide resolution at the binomial scale (avoidance
or no avoidance). Encounter duration has previously been
used to quantify avoidance by Hectors dolphins
Cephalorhynchus hectori of swimmers in New Zealand
(Bejder et al., ) and to assess long-term trends in en-
counters with dwarf minke whales Balaenoptera acutoros-
trata on the Great Barrier Reef, Australia (Birtles et al.,
). However, these studies used different definitions of
encounter duration: Bejder et al. () defined it as the
length of time that a swimmer spent within  mof
the nearest dolphin and Birtles et al. () defined it as
the time between the first sighting of a whale and the end
of the vessels contact with that whale. Here we define
encounter duration as the time between the first swimmer
entering the water in the presence of a whale shark and
the last swimmer returning to the boat. This definition
could be less useful in areas with significant currents or
where the boat would take longer to retrieve swimmers.
We found a positive relationship between the number of
swimmers and encounter duration. This result may be an
artefact of the definition in use, as larger groups of swim-
mers are likely to contain members with a wider range of
swimming abilities. The presence of more able swimmers
is likely to lead to an increase in encounter duration, as
they are able to maintain visual contact with a shark for
longer than slower or less fit individuals. Another important
point to consider in future work is the number of swimmers
that are actually in contact with the shark from a behav-
ioural perspective, investigating how swimmer proximity
Tourism and whale sharks 5
©2014 Fauna & Flora International,
Oryx
,18
may influence these results. Future studies that rely on in-
water observers could trial re-defining encounter duration
as the time between the observers first and last visual con-
tact of the shark underwater, to reduce the influence of
swimming ability and group size on results.
Consistent with results from Quiros () but contrary
to Norman () encounter duration was significantly
longer with scarred whale sharks. This could be because
scarred individuals have slower reaction times and a re-
duced level of agility as a consequence of their previous
injuries. However, the severity of scarring did not signifi-
cantly alter encounter duration. Another hypothesis is that
older individuals, who would presumably be more likely to
have accumulated scars, show less avoidance than younger
individuals. However, we found no significant relationship
between shark size and the severity of scarring. A third
plausible explanation is that some sharks have a slower nat-
ural reaction time to potential threats and are therefore
more likely to accumulate injuries than sharks with faster re-
actions. Sharks may also vary in their inquisitiveness.
Further research is required to confirm or refute these
hypotheses.
A mean of .swimmers per encounter was recorded.
This exceeds the maximum recommended by most manage-
ment plans, which limit swimmer numbers to three in
Mexico (Remolina Suárez et al., ) and  in Australia
(Catlin & Jones, ), although these limits may be par-
tially influenced by operational constraints in those loca-
tions. From a management perspective it is also important
to consider customer satisfaction with the experience, which
can be significantly reduced in crowded scenarios (Davis
et al., ).
The inclusion of well-defined and easily observed para-
meters of known significance that can be recorded by either
trained specialists (e.g. researchers) or non-specialist obser-
vers (e.g. tour operators or volunteers) would increase the
amount of information available, enabling routine sustain-
ability assessments and facilitating inter-site comparisons
(Graham, ). We have created a template data collection
sheet (Supplementary Material ), which is currently in use
in Mozambique. We also encourage the submission of
standardized photographs of all whale sharks encountered
to the global whale shark photo-identification library
(Wild Me, ). This facilitates the incorporation of
individual-based analyses into future studies, concurrently
enabling more accurate regional and ocean-wide studies of
population size and interconnectivity (Graham, ;
Brooks et al., ).
With increasing global interest in human interactions
with whale sharks there is a clear need for the implementa-
tion of monitoring initiatives at tourism sites to understand
and avoid long-term anthropogenic effects. Other import-
ant whale shark tourism sites, such as those in Mexico,
the Seychelles and Western Australia, have similar
characteristics in terms of population structure. The results
of this study demonstrate that encounter duration is a useful
metric for assessing whale shark behaviour, and provide a
template for future studies in Mozambique and other
locations.
Acknowledgements
We thank the All Out Africa volunteers and staff for their
efforts in collecting the majority of the data used in this
study, and Marine Megafauna Foundation staff, volunteers
and students who also contributed. We thank the developers
of, and contributors to, the Wildbook whale shark database.
The support of Casa Barry Lodge and Tofo Scuba through-
out the field portion of the project is greatly appreciated.
This study was made possible by support from the
National Marine Aquarium (UK), Shark Foundation, GLC
Charitable Trust, Project AWARE Foundation, the Rufford
Small Grant Foundation, PADI Foundation, Ocean
Revolution, Fondation Ensemble and private donors. We
thank Philip Dearden, Adrian Gutteridge, Conrad Speed
and the anonymous reviewers for their comments on this
article.
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Biographical sketches
PETER HASKELL specializes in behavioural ecology and is interested
in the role that tourism can play in elasmobranch conservation.
ANDREW MCGOWANSresearch interests include avian cooperative
breeding, conservation of sea turtles, and tropical seabird populations.
ANNA WESTLING is a coral reef ecologist interested in citizen science.
ADRIANA MÉNDEZ-JIMÉNEZ is interested in the h uman dimension of
the environment, and in the way people perceive environmental is-
sues. CHRISTOPH ROHNER is interested in the movement and feeding
ecology of large pelagic animals, and linking their behaviour to bio-
logical and oceanographic drivers. KYM COLLINS specializes in the
behaviour and acoustics of marine mammals. MARCELA ROSERO-
CAICEDO is interested in marine conservation and specializes in sci-
entific and environmental communication in volunteer programmes
and local communities. JODI SALMOND studies Australian reef eco-
systems in a global context, specializing in citizen science and marine
education. ARA MONADJEM is interested in the application of quan-
titative ecological techniques to conservation-related problems in
small mammals and birds. ANDREA MARSHALL studies the biology,
ecology, monitoring and management of manta rays globally. SIMON
PIERCE is a marine conservation ecologist specializing in whale
sharks.
8 P. J. Haskell et al.
©2014 Fauna & Flora International,
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,18
... Harpy Eagle tourism can help to generate concrete financial value for habitat conservation, as has happened with other predators (Macdonald et al. 2017, Tortato et al. 2017. Although practical protocols have been developed for better practices of wildlife tourism (Haskell et al. 2015), few have addressed Amazonian wildlife. Creating evidence-based visitation schedules and tailoring them to offer higher viewing probabilities can help jumpstart the region's potential. ...
... Those discoveries are of prime interest regarding Harpy Eagles as a wildlife attraction and the management of tourism that fits nest visitation schedules of eagles. Responsible and controlled schedules for tourists viewing nests are particularly important for sensitive species with multi-decadal life cycles, low breeding potential, and high tourism value (Ashe et al. 2010, Haskell et al. 2015, Tortato et al. 2017. For Harpy Eagles, our data showed that both adults and eaglets were mainly active during the early and late hours of the day, with a higher peak in activity during the morning (10h15-10h45) for adults and late afternoon (15h15-15h45) for fledgling, adult-sized eaglets. ...
... The operation of responsible, sustainable, profitable Harpy Eagle nest photo-tourism is a laborious and expensive process, as towers must be moved, and new agreements reached with landowners. Besides threatening wildlife, poor management decisions would ultimately affect the attractions they were built on, compromising the sustainability of the business model (Haskell et al. 2015). Furthermore, having Harpy Eagle nests under a relatively constant watch could open the door to several avenues of conservation actions in the face of fairly heavy deforestation. ...
Article
Full-text available
Tourism can be a powerful tool for wildlife conservation if well controlled and responsibly managed. Apex predators constitute particularly attractive subjects for tourism, but simultaneously they may generate conflict with local communities. Harpy Eagles Harpia harpyja are the largest eagle species and are highly sought-after by ecotourists. The last stronghold of the Harpy Eagle is the Amazon Forest, which is being deforested for cattle ranching. We tested methods for developing Harpy Eagle ecotourism as a potential tool to harmonize these issues. Using camera traps, we collected data on timing of Harpy Eagle visits to their nests, as well as on probabilities of viewing an eagle. Harpy Eagles can only be seen predictably during the first 12 of the 30–36 month nest cycle. In nests with nestlings (up to 5–7 months), adults are visible on a daily basis, and this period lasts 16.6% of the nesting cycle, demanding a minimum of 13, 17, and 26 nests to have at least one nest with a nestling on 90%, 95% and 99% of the days. After this 5–7 month window, we found that two and 4.16 days spent at nests afforded high probabilities of sighting a fledgling or adult eagle, respectively. Harpy Eagles were mainly active at the beginning and the end of the day. Activity core lasted 6.5 decimal hours for adults, peaking at 10h00, and 7.45 decimal hours for fledged eagles, peaking at 15h00. Our results demonstrate that Harpy Eagles fit several criteria for a viable wildlife attraction: predictable in activity and location, viewable, and diurnal, even though at the same time they are considered a rarity. In a broader perspective, Harpy Eagle tourism shows every indication of being a significant tool for more robust rainforest conservation.
... A similar pattern was observed in Mozambique. Pierce et al. (2010) found that 34.6% of observed encounters between an individual shark and swimmers resulted in avoidance responses at Tofo Beach, Mozambique, while Haskell et al. (2014) found that 64.7% of observed interactions resulted in avoidance responses at the same site suggesting that the sharks' tolerance for tourism activities also decreased over the study period. ...
... Furthermore, Pierce et al. (2010) and Haskell et al. (2014) found that sharks exhibiting avoidance behaviors resulted in shorter encounters indicating that tourist presence did affect the shark's short-term behavior. However, Pierce et al. (2010) noted that encounter length was also dependent on the shark's prior reaction to the boat, such that in-water interactions with sharks displaying boat avoidance behaviors were significantly shorter than interactions with sharks that did not display such avoidance behaviors. ...
... Insert Table 10.4 here Insert Table 10.5 here There are, however, conflicting results regarding the effect of swimmer presence on whale sharks. For example, Araujo et al. (2017) and Haskell et al. (2014) found that the number of previous encounters with tourists a shark experienced did not affect interaction duration in Leyte and Mozambique. Quiros (2007), meanwhile, found that sharks that had been swum with repeatedly were more likely to exhibit a dive response compared to those that had been sighted for the first time, which suggests that sharks learn tourist avoidance over time. ...
... Increased drag from carrying fishing gear can incur considerable energetic costs (Moore and Van der Hoop, 2012;Van der Hoop et al., 2016), while severe tissue damage can result in haemorrhage or debilitation (Cassoff et al., 2011;Moore and Van der Hoop, 2012), and open, unresolved wounds can lead to serious infection (Borucinska et al., 2002;Cassoff et al., 2011). Injured whale sharks (Rhincodon typus) exhibit less evasive behaviours toward tourists and boats than non-injured individuals (Quiros, 2007;Haskell et al., 2015), which suggests that injuries may reduce their agility, or that individuals are choosing warmer surface waters, where tourism activities are concentrated, to aid wound healing (Womersley et al., 2021). Injuries have also been found to disrupt social behaviours of bottlenose dolphins (Tursiops truncatus), which may compromise their long-term survival, or make them more vulnerable to predation (Greenfield et al., 2021). ...
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Manta ray populations worldwide are vulnerable to sublethal injuries resulting from human activities, e.g., entanglement in fishing line and boat strikes, which have the potential to impact an individual’s health, fitness, and behaviour. Sublethal injuries and physical abnormalities also occur naturally from predation events, deformity, parasites, and disease. To determine the type and frequency of anthropogenic and natural originated injury events affecting Mobula alfredi and M. birostris in the Maldives, we examined data from the Manta Trust’s Maldivian Manta Ray Project (MMRP) database, which contains 73,638 photo-identification (photo-ID) sightings of the two manta ray species from 1987 to 2019. The likely origin of each injury or physical abnormality was determined based on visual assessment of the photo-ID images. Multiple injuries to an individual originating from the same event were grouped for analysis. Generalised linear mixed models (GLMM) were used to investigate the relationship between the occurrence of injury events and the explanatory variables sex and maturity status for both species, with the additional variable site function (cleaning, feeding, cruising) investigated for M. alfredi. Spatial and temporal variations in M. alfredi injury events, and their origin and type, were investigated by calculating the percentage of injury events per sighted individual at each Maldivian atoll, and per re-sighted individual in each year from 2005 to 2019. For both species, injury events were predominantly of natural origin, with predatory bites being the most frequent type. The most common anthropogenic injury type was entanglement in fishing line. Injuries to M. alfredi were significantly more likely to be observed on juveniles than adults, males than females, and at cleaning stations as opposed to feeding or cruising sites. Neither sex nor maturity status were significant explanatory variables for the occurrence of injuries to M. birostris. Highest percentages of anthropogenic injuries per sighted M. alfredi were recorded in North Malé, South Malé, Baa, Addu, and Laamu Atolls, where boat traffic, fishing, and tourism activities are concentrated. Overall, this work greatly improves understanding of the sublethal threats faced by manta rays in the Maldives; identifying focus areas where conservation management actions are required to ensure more effective protection of this threatened species group.
... 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. ...
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• 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 ...
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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.
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Growth rings (GR) in vertebral centra of 15 whale sharks, Rhincodon typus, four female (418–750 cm precaudal length), 10 male (422–770 cm), and one of unknown sex (688 cm), were examined using x-radiography. GR counts were made from scanned images and count precision was determined using the average percentage error index (4.19%) and the index of precision D (3.31%). In females, counts ranged from 19 GR (418 cm) to 27 GR (750 cm); in males from 20 GR (670 cm) to 31 GR (770 cm). Three mature males had 20 GR (670 cm), 24 GR (744 cm) and 27 GR (755 cm). A female with 22 GR (445 cm) was adolescent. There was a linear relationship between centrum dorsal diameter and body length, and back-calculated body lengths at number of GR are presented. A linear relationship between body length and number of GR prevented the calculation of von Bertalanffy parameters from either observed or back-calculated values.
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Most arguments invoked so far by the scientific community in favour of shark conservation rely on the ecological importance of sharks, and have little impact on management policies. During a 57-month study, we were able to individually recognise 39 sicklefin lemon sharks that support a shark-feeding ecotourism activity in Moorea Island, French Polynesia. We calculated the direct global revenue generated by the provisioning site, based on the expenses of local and international divers. The total yearly revenue was around USD5.4 million and the 13 sharks most often observed at the site had an average contribution each of around USD316 699. Any one of these sharks represents a potential contribution of USD2.64 million during its life span. We argue that publicising economic values per individual will be more effective than general declarations about their ecological importance for convincing policy makers and fishers that a live shark is more valuable than a dead shark for the local economy. Studies monitoring the potential negative ecological effects of long-term feeding of sharks should, however, be conducted to ensure these are also considered. Besides declarations about the non-consumptive direct-use value of sharks, as promoted by ecotourism, the calculation of their other economic values should also benefit shark conservation.
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The Western Australian Department of Environment and Conservation has statutory responsibility for conservation, and commercial and recreational aspects of the Ningaloo Whale Shark Experience. The whale shark experience participation rate had grown by 150% since 1995. In this paper, the ability of the existing management programme to balance ecological, social and economic factors is reviewed. The Department has established a statutory Code of Conduct which applied equally to commercial and recreational users when interacting with whale sharks. The Code of Conduct for Whale Shark Interaction was developed to provide for an acceptable impact on the wildlife and a safe visitor experience. A restricted licensing regime was applied. Studies conducted to date indicate that any significant impact on the whale shark population is unlikely as a result of the tourism interactions at Ningaloo. Social surveys indicate a high level of satisfaction with all aspects of the whale shark experience. An education programme was implemented to cater for industry needs and a local, national and international audience. At this time, the Ningaloo Whale Shark experience may effectively be described as an ecologically sustainable wildlife tourism industry managed by a government conservation department based on non-consumptive use of wildlife.