Integrating citizen science and telemetry techniques in understanding the movement patterns of the
whale shark (Rhincodon typus)
Bradley Michael Norman
This thesis is presented for the Doctor of Philosophy
2016
Murdoch University, Perth, Western Australia
1
DECLARATION
I declare that this thesis is my own account of my research and contains as its main content work which has not previously been submitted for a degree at any tertiary education
institution.
I wish to acknowledge and thank the following researchers for data submitted to the ECOCEAN / Wildbook Library and their input to Chapter 2 including: Jason Holmberg, Zaven Arzoumanian, Samantha Reynolds, Rory Wilson, Dani Rob, Simon Pierce, Adrian Gleiss, Rafael de la Parra, Beatriz Galvan, Deni Ramirez-Macias, David Robinson, Steve
Fox, Rachel Graham, David Rowat, Matthew Potenski, Marie Levine, Jennifer A. McKinney, Eric Hoffmayer, Alistair Dove, Robert Hueter, Allesandro Ponzo, Gonzalo Araujo, Elson Aca, David David, Richard Rees, Alan Duncan, Christoph Rohner, Claire Prebble, Alex
Hearn, David Acuna, Michael Berumen, Abraham Vázquez, Jonathan Green, Steffen Bach, Jennifer Schmidt, Stephen Beatty, George Shedrawi, Steve Lindfield, Darcy Bradley, Lisa
West and David L. Morgan. In addition, this research has made use of data and software tools provided by Wildbook for Whale Sharks, an online mark-recapture database operated by the non-profit scientific organization Wild Me with support from public donations and the Qatar
Whale Shark Research Project.
……………………………………………………………………………………...
Bradley Michael Norman
2
Acknowledgements
The completion of my studies for this thesis was only achievable with the support of a fantastic yet diverse group of friends and colleagues. First and foremost, I’d like to sincerely thank my Principal Supervisor Dr David Morgan for his advice throughout the project and his belief in my work as a whole. A very big thank you also to other Murdoch University staff, including Professor Neil Loneragan for his input and assistance, especially in the early stages of the study. Thanks also to Murdoch University for providing my PhD Scholarship (MURS).
I am indebted to the vision and action of two of the smartest people I have had the pleasure to work with, namely Jason Holmberg and Zaven Arzoumanian who really helped put the Whale Shark Library front and centre for the global monitoring of this species.
ECOCEAN Inc. has been pivotal throughout and enabled my work to extend to community education and engagement and I am extremely grateful for this opportunity. Through the Rolex Award for Enterprise and National Geographic Society, it has been possible to expand the project to a global stage.
The Ningaloo community of Whale Shark Ecotourism Industry stakeholders have always supported my research and helped whenever the request went out – especially with space on commercial vessels to undertake the photo-identification research. They have provided data and whale shark identification images that have helped make the project the most comprehensive monitoring program on this species worldwide. This support has also been very much appreciated from staff and friends at the Western Australian Department of Parks and Wildlife who continue to facilitate my research on the whale sharks at Ningaloo.
Fieldwork at Ningaloo has been supported over the years by many stakeholders for which I am sincerely grateful, including Novotel Ningaloo Resort, Exmouth Cape Tourist Park, Lighthouse Caravan Park, Potshot Hotel Resort, IGA Exmouth, Steve Wall, Alex Kailis (MG Kailis), Olympus, Cressi, Tusa and Mares.
This long-term study has received input, advice and an enormous amount of help from friends, colleagues, passionate volunteers and members of the general public. In no particular order, I’d like to acknowledge each as central to the ultimate successful completion of this massive research project, including: Professor Rory Wilson, Professor Lyn Beazley, Dr Nikolai Liebsch, Dr Adrian Gleiss, Dr Stephen Beatty, Jeff Whitty, Mark Allen, CSIRO (Dr John Stevens, Dr Barry Bruce, Dr Russ Babcock), AATAMS, OzTrack, CLS-Argos (Guan Oon and Holly Laurie), IVEC, George Shedrawi, Steve Lindfield, Darcy Bradley, Kathy Zischka, Helen Shortland-Jones, James Catlin, Lisa West, Hester Bushell, Emily Hamley, Aleisha Caruso, Darren Cossill, Heath Bentley, Jennifer Spark, Justin Palmer, Christophe Baudia, Caroline Confort, Sophie Bernstein, Aisleen Dilks, Taylor Mathews, Eden Baxter, Briana Canny, Andrew Needham, Kim Hands, Kylie Maguire, Cameron O’Neill, Kaitlyn
3
Close, Dennis Beros, Peter and Heather Lake, Exmouth District High School, Earthwatch Institute and volunteers, and Mark Norman.
Securing the funding required to undertake this extensive study has always been a challenge, but I have had some outstanding supporters throughout, including: Steve Wall, Luc Longley, Anna Gare, Tim and Megan Caporn, Warwick Mathews, Steve Perry, Jock Clough, Tim Winton, Kelly Slater, John Butler, Xavier Rudd, Buzz Aldrin, Brian Eno, Sir David Attenborough, Ron and Valerie Taylor, Shelley Taylor-Smith, Paul Watson, Olivia Samec, Leon Holmes, Glen Cowans, Beach House Art, Peter Rigby, Rosendorff Jewellers, Driftwood Jewellers, Smales Jewellers, Watches of Switzerland, Hearts for Sharks, Tourism WA, Graeme Meinema (Fisheries WA), and Rolex Australia.
In addition, small grants have been forthcoming from the following, and I am extremely appreciative for this assistance: Thyne Reid Education Trust, Winifred Violet Scott Estate, Garry White Foundation, Holsworth Wildlife Research Endowment, Australian Geographic Society, The Tony and Lisette Lewis Foundation, Perth Convention Bureau, Australian Government Community Action Grants, Coast and Clean Seas, Coastwest/Coastcare, HSBC Bank, Australian Marine Conservation Society, United Nations Environment Programme (UNEP), Western Australian Natural Resource Management grants.
Last but definitely not least, I wish to say a huge thanks to my folks Lyn and Alan for their ongoing support from the very outset, and especially to Samantha Reynolds for her love, assistance and the final drive to the finish line!
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Abstract
Since its inception in 1995, the whale shark photo-identification library that was developed
for whale sharks at Ningaloo Reef (Western Australia) has burgeoned, with submissions from
over 54 countries throughout the species global range and it has received more than 30000
encounter reports with sighting information and associated photographic images. In recent
times, with the assistance of digital techniques, a global campaign to promote the library, and
an increase in the number research active organisations utilising the library, has led not only
to the identity of the world’s 20 ‘hotspots’ for whale shark aggregations, but to improved
conservation measures for this internationally threatened species.
The collation and analysis of such data has demonstrated the uniqueness of the
different ‘population’ centres, inter-jurisdiction movement patterns within only adjacent sites,
an absence of oceanic basin migrations, and has implied that many of the aggregations are
comprised of individuals that show a strong affinity to that area, and some individuals return
for decades. The population demographics of the various sites may be either homogenous or
heterogeneous, including some sites which are seasonally used, while others have year-round
residents. Some populations are dominated by juvenile males, while others may be entirely
large females. Globally, more than 6000 individual whale sharks have been identified, with
over 1000 identified from Ningaloo Reef, a location where contemporary belief was that the
aggregation was extremely seasonal, occurring during the Austral autumn and winter. More
than 8000 useable images of whale sharks were analysed from Western Australia, and a
targeted campaign further assessed whale shark occurrences from 3000 km of the Western
Australian coastline. This, coupled with satellite tracking and acoustic tracking, identified
that not only are whale sharks found at Ningaloo throughout all months of the year, but the
species within Western Australian waters has a strong site fidelity to Ningaloo during the
main aggregation period, although individuals may roam north, west or south from Ningaloo
and the species ultimately spans much of the State’s immense coast. Acoustic tracking
further revealed that at Ningaloo, individuals remain relatively close to the outer reef edge,
moving longitudinally. Satellite tracking and citizen science allowed the hypothesis to be
tested that in Western Australian waters, individuals may be seeking other areas of high
5
productivity outside of the predictable mass coral spawn at Ningaloo Reef, which has long
been thought to be a significant driver of their build up in numbers during the Austral
autumn. A number of the whale sharks photo-tagged in Western Australia have been ‘re-
captured’ many times over a 21 year period (1995-2015), and these individuals have been
studied at a time-scale that exceeds almost all other studies of individual fish. The
continuation of the monitoring program of this long-lived species, together with the
continued update of citizen science in the region, will in time reveal and help solve additional
mysteries (including migration patterns and critical habitats) surrounding this enigmatic
species.
6
Table of Contents
Acknowledgements…………………………………………………...….. 3
Abstract……………………...……………………………………….…... 5
Chapter 1
General Introduction…………………………..……………………..….… 8
Chapter 2
Undersea constellations: ‘Citizen Scientists’ elucidate the global biology of a threatened marine mega-vertebrate…………………….……………..… 21
Chapter 3
The efficacy of photo-identification monitoring whale sharks at Ningaloo Marine Park, Western Australia…………………..……….…………….… 46
Chapter 4
Residency of whale sharks at Ningaloo Marine Park informed using acoustic telemetry and photo-identification……….…………………………….….. 68
Chapter 5
Does the whale shark aggregate along the Western Australian coastline beyond Ningaloo Reef?.............................................................................................. 88
Chapter 6
General conclusions……...……………………..…………….................… 113
References………………………………………………………..….……. 118
Appendix 1……………………………………...………………..….….… 134
Appendix 2……………………………………...……….…………..….… 195
Appendix 3……………………………………...……….…………..….… 200
Appendix 4……………………………………...……….…………..……. 204
7
Chapter 1 General Introduction
The whale shark (Rhincodon typus) is not only the largest fish, it is the largest non-cetacean
animal in the world, reaching over 18 m in total length and weighing over 20 tonnes (Chen et
al. 1997, Compagno 2001, Borrell et al. 2011). The common name “whale shark” is derived
from the fish’s morphology, being as large as many whales and also a filter feeder like many
baleen whale species. It is one of only three known filter feeding shark species (along with
the basking shark (Cetorhinus maximus) and the megamouth shark (Megachasma pelagios)
(Last and Stevens 1994). Whale sharks feed predominantly on zooplankton, including red
crab larvae, copepods, tropical krill and small fish (Norman 1999).
Primarily pelagic, R. typus inhabits all tropical and warm-temperate seas (Compagno
2001, Last and Stevens 1994). Individuals of this species have a brown or bluish-grey body
with a white belly (Last and Stevens 1994). The skin, which can be in excess of 10 cm thick
(Gifford et al. 2007), is marked with a checkerboard pattern of white spots and stripes that are
unique to each individual, making it an ideal species where photo-identification can be used
(Taylor 1994, Norman 1999, Arzoumanian et al. 2005, Van Tienhoven et al. 2007).
Whale sharks have been found to migrate great distances and annual feeding
aggregations are seen at various places around the world, including Ningaloo Reef (Western
Australia), Belize, off the Yucatan Peninsula in Mexico, Seychelles and near Donsol in the
Philippines (Gifford et al. 2007, Graham and Roberts 2007, Holmberg et al. 2008, 2009, de la
Parra et al. 2011). The Ningaloo Reef aggregation is believed to be one of the most
predictable and occurs after the mass spawning of corals in March and April (Norman 1999).
This species may also migrate to breed (Sequeira et al. 2013), although neither mating
nor pupping has been observed. The capture of a pregnant female R. typus in July 1996 off
8
Taiwan with approximately 300 pups demonstrated that whale sharks are obligate
lecithotrophic livebearers (Joung et al. 1996). There is evidence that the pups are not all born
at once, but rather that the female retains sperm from one mating event and disperses pups
over a prolonged period (Schmidt et al. 2010).
Hsu et al. (2014a) suggest an age and total length (TL) at sexual maturity for female
R. typus is 19-22 years (between 8.7 and 9.6 m TL), while 50% of males reach sexual
maturity at 8.1 m TL (Norman and Stevens 2007) or at about 17 years of age (Hsu et al.
2014a).
A recent review of available data on whale shark sightings tracked movements and
distribution information (see Sequeira et al. 2013), providing some evidence for the
hypothesis of broad-scale connectivity among populations throughout the world’s oceans.
This further supports aspects of previous DNA studies undertaken by Castro et al. (2007) and
Schmidt et al. (2009).
The whale shark is listed as ‘vulnerable to extinction’ on the International Union for
the Conservation of Nature (IUCN) Red List of Threatened Species (Norman 2000). It is also
listed on Appendix II of the Convention on International trade in Endangered Species
(CITES) and Appendix II on the Bonn Convention for the Conservation of Migratory Species
(CMS). Although now protected in some countries, the migratory nature of the threatened R.
typus may result in their moving from a protected area to an area where they may be hunted
(Bradshaw et al. 2008, Li et al. 2012, Akhilesh et al. 2013). Fishing pressure has been
implicated in population decline for this species (Pierce and Norman 2016). Human induced
habitat destruction is also a major threat to this filter-feeder (Pierce and Norman 2016), which
is dependent upon food pulses and critical habitats to survive. The global population of this
threatened species remains unknown, although research is being undertaken using various
9
techniques at locations throughout the world in an attempt to shed light on this question (see
Rowat and Brooks 2012).
Wildlife monitoring options
Wildlife monitoring can take many forms depending on the species, the environment in
which it is used, and the type of data required (Baratchi et al. 2013). Fauna surveys for
example, can employ non-invasive methods such as aerial surveying, camera traps, scat
detection/analysis, and direct observation (Garden et al. 2007, De Bondi et al. 2010).
Alternatively, animals can be physically tagged (with markers to facilitate monitoring and
avoid duplication when counts are taken) and/or fitted with electronic monitoring devices.
These types of invasive tagging can however result in some level of stress to the individuals
in question (Wilson 2011).
Genetic sampling
Genetic studies generally require a tissue sample to be collected for subsequent analysis
(Schmidt et al. 2009, 2010), although with cetaceans for example, skin sloughed off and
available in the water near an animal (Amos et al. 1992) can be gathered without impacting
the animal. Similarly, faeces, shed hairs, or shed feathers can be collected from wild animals
as a source of DNA (Taberlet and Luikart 1999, Waits and Paetkau 2005).
For whale sharks however, tissue samples are most commonly collected using a
biopsy dart to penetrate the skin of the animal to remove the required piece of organic
material (Schmidt et al. 2009).
10
Marker tags
Numbered tags can be used to aid in confirmation of a resighted individuals for population
and distribution studies including e.g. small sawfish (Pristis pectinata Latham)
(Simpfendorfer et al. 2008), various bird species (Miller‐Rushing et al. 2008), and even
Quokkas (Setonix brachyurus) (Shield and Woolley 1961). When used on whale sharks,
marker tags can impact the animal when first deployed as these are secured to the animal via
a tag ‘anchor’ that must penetrate the skin (which can be 10 – 15 cm thick (Gifford et al.
2007)). This technique can be used to identify individuals at or returning to a particular area
(Hueter et al. 2013). The tags are however known to often break and become covered with
fouling (Graham and Roberts 2007, Bansemer and Bennett 2008), thereby rendering them of
questionable value for monitoring purposes.
Biotelemetry and biologging
While both telemetry and logging involve the remote monitoring of animal movements,
behaviour, physiology, and/or environmental information, the means of collection are
different: e.g. with telemetry, a signal from an animal-borne transmitter is sent to a receiver,
while with logging, information is recorded and stored in archival logger which is carried by
the animal, and these data are downloaded when the device is retrieved (Myers et al. 2006,
Cooke 2008, Rutz and Hays 2011).
Several types of electronic tags have been used in research on R. typus, with each
focussing on the collection of important yet different data (Gunn et al. 1999, Graham and
Roberts 2007, Sleeman et al. 2010). While satellite tags provide data on the movements of
whale sharks by providing accurate position of the shark, these must have an uplink to
orbiting satellites (Acuna-Marrero et al. 2014). Acoustic tags require a listening station to
receive data transmitted through water (Cagua et al. 2015) while archival tags must be
11
retrieved in order to download the collected data (Gleiss et al. 2011b). In each case, the
animal must either be restrained to deploy the tag/s or the tags attached via a number of
methods, all of which requires contact with the animal in question (Hsu et al. 2007, Gleiss et
al. 2009).
Acoustic telemetry
Acoustic telemetry for fish research employs acoustic tags, where small sound-emitting
devices are deployed allowing the detection and/or remote tracking of fish in three
dimensions (Pincock et al. 2010). Acoustic telemetry (including acoustic tags) can therefore
be commonly used to monitor the behaviour of fish. Acoustic tags transmit a sound signal or
"ping" that sends location information about the tagged fish to the hydrophone receiver. By
tying the received acoustic signature to the known type of programmed signal code, a specific
fish is identified (Heupel and Hueter 2001, Heupel et al. 2006).
Acoustic studies have been undertaken on species such as Spangled
Emperor (Lethrinus nebulosus) (Pillans et al. 2014), the Caribbean reef shark (Carcharhinus
perezi) (Bond et al. 2012), and the giant manta ray (Manta birostris) (Dewar et al. 2008)
revealing new insights to habitat use, periods of residency, and movement patterns. Whale
sharks have also been studied in this way (Cagua et al. 2015).
Archival data loggers
An archival tag is a data logger that uses sensors to record data at predetermined intervals
(Sibert 2001). Data storage tags usually have a large memory size and a long lifetime, and
most archival tags are supported by batteries that allow the tag to record positions for several
years (Arnold and Dewar 2001). Archival tags can have a variety of sensors; temperature,
depth, light, salinity, pressure, pitch and roll, GPS, magnetic and compass (Gunn and Block
12
2001, Whitney et al. 2012). They can be used internally or externally (Bridger and Booth
2003), have been used on a wide variety of marine and terrestrial animals from yellowfin
tuna (Thunnus albacares) (Schaefer et al. 2007) to sooty shearwaters (Puffinus
griseus) (Shaffer et al. 2006). This technology has also been deployed on whale sharks
(Wilson et al. 2006, Hueter et al. 2013).
The Daily Diary is an archival data-logging tag that has two sensory systems that help
determine behaviour directly via change in orientation or movement, these being the tri-axial
magnetometer (compass) and tri-axial accelerometer (Wilson et al. 2008). The accelerometer
is a small electronic device used to monitor changes in an animal’s acceleration and
simultaneously measures along all three axes i.e. forwards, up/down, and sideways. The
Daily Diary is also able to determine the animal’s speed, direction and position (Wilson et al.
2008). Species on which these tags have been deployed are as diverse as the Andean condor
(Vultur gryphus) (Shepard et al. 2011), the northern fur seal (Callorhinus ursinus) (Benoit-
Bird et al. 2013) and the dromedary camel (Camelus dromedaries) (Bidder et al. 2014). Daily
Diaries have also been used on whale sharks at Ningaloo (Gleiss et al. 2009, 2011b, 2013).
Satellite tags
Satellite telemetry can be used to collect and transmit movement, behaviour and
environmental data from a tag deployed on free-ranging animals, and does not require manual
retrieval of the tag (Patterson and Hartmann 2011). Pop-up archival transmitting (PAT) tags
collect, store and ultimately transmit these data to overhead satellites when the tag detaches
and floats to the surface, enabling a snapshot of the horizontal movements and diving
behaviour of marine animals (Davis et al. 2007).’ These tags have been used on species
ranging from tuna (Thunnus thynnus) (Block et al. 2005), to sunfish (Mola mola) (Sims et al.
13
2009), to jumbo squid (Dosidicus gigas) (Gilly et al. 2006), to reef manta rays (Manta
alfredi) (Braun et al. 2014).
A Platform Terminal Transmitter (or PTT) is a small satellite transmitter and when
used in marine applications, transmits a signal which is received by overhead satellites when
free of the water, enabling an accurate position fix to be obtained. These units have been used
on a variety of species ranging from turtles (Dermochelys coriacea) (Hughes et al. 1998) to
sea lions (Neophoca cinerea) (Fowler et al. 2007) to killer whales (Orcinus orca) (Durban
and Pitman 2011).
For shark studies, PTT’s can be either embedded in positively buoyant syntactic foam
torpedo-shaped floats and attached via a braided stainless steel tether to a surgical grade
stainless steel tag anchor deployed on the body of a shark (Sleeman et al. 2010, Hearn et al.
2013), or can be attached to the 1st dorsal fin via bolts and nuts (Hammerschlag et al. 2011,
Fitzpatrick et al. 2012). The technology utilises a satellite-based system (ARGOS) which
receives transmissions from the tags and enables calculation of the animal’s geolocation in
successive uplinks (Costa et al. 2012). ARGOS has the unique ability to geographically
locate the source of the data (i.e. the tagged shark) anywhere throughout the globe, without
the need to retrieve the tag. In order to transmit to the overhead satellite however, the tag’s
antenna must be free of the water surface at the same time orbiting satellites are passing
overhead (Hammerschlag et al. 2011).
On-animal video
National Geographic ‘Crittercam’ (Marshall 1998) and Customised Animal Tracking
Solutions ‘CATScam’ have been developed to record video from an animal (e.g. tiger shark
(Galeocerdo cuvier), white shark (Carcharodon carcharias)) as it swims (at the surface and
throughout the water column), and then release to float to the surface where it is located
14
(using the VHF transmitter embedded in the unit) and subsequently retrieved (Heithaus et al.
2001, Chapple et al. 2015). This enables the behaviour of the tagged animal to be recorded
without the influence of human presence, enabling researchers to ‘see’ what other species and
varied conditions a tagged animal may encounter.
Photo-identification
An important requirement in many monitoring programs is to minimise the effect of the
observer on the observed. Non-invasive tagging can facilitate this. Using natural markings
on the skin of an animal to identify individuals has been used successfully for many species
of terrestrial animals (Petersen 1972, Kelly 2001, Jackson et al. 2006) and marine species
(e.g. humpback whales (Megaptera novaeangliae) (Mizroch et al. 1990), orcas (O. orca)
(Young et al. 2011), bottlenose dolphins (Tursiops truncatus) (Markowitz et al. 2003) and
grey nurse sharks (Carcharias taurus) (Van Tienhoven et al. 2007). Whale sharks are an
excellent example of another species where this has provided important information on the
residency and resighting rates of individuals at Ningaloo Reef and other locations throughout
their range (Norman 1999, Arzoumanian et al. 2005, Holmberg et al. 2008, 2009, Rowat and
Brooks 2012).
Many elasmobranch species may be suitable for photo-identification monitoring
because of spots/pigmentation patterns and accessibility to the tourism industry, including
great white sharks (C. carcharias), manta rays (Manta birostris), zebra sharks (Stegostoma
fasciatum), and tiger sharks (G. cuvier) (Bansemer and Bennett 2008). Whale sharks too are
born with unique body pigmentation (Norman 2004). This natural patterning of lines and
spots shows no evidence of significant change over years and may therefore be used to
identify individual sharks (Taylor 1994, Norman 1999): its uniqueness has been corroborated
by traditional tagging and identifications made based on scarring and other visual markers
15
(Taylor 1994, Norman 1999, Arzoumanian et al. 2005). Through the combination of
photographed encounters and spot-pattern matching, a whale shark may be ‘tagged’ without
physical contact or interference with the animal.
Citizen science
Citizen science involves volunteers in research (Dickinson et al. 2010). Millions of people
(often non-scientists) are now being engaged in such citizen science projects worldwide to
collect, catalogue and assist in the analysis of scientific data, thanks in part to the increase in
the number of web-based projects available (Parsons et al. 2011, Bonney et al. 2014,
Theobald et al. 2015). Through the engagement of citizen scientists, new insights as diverse
as the rediscovery of possibly extinct butterfly species (Lawrence 2015); shifts in the timing
of crucial events in plant life cycles in response to climate change (Gonsamo et al. 2013),
mapping of species distributions and migration routes (Lees and Martin 2014, Pocock et al.
2015, van der Wal et al. 2015); and even an assessment of anthropogenic modifications of
Earth's biosphere (Kyba et al. 2013) have been possible.
The precursor for a global citizen science monitoring project for whale sharks was
initiated in 1995 with the development of a photo-identification library at Ningaloo Reef,
Western Australia using photographs of individual sharks examined for identifying
characteristics, including spot patterns (see Norman 1999). To support the collection and
centralisation of biological data by wildlife researchers, the Shepherd Project began in 2002
with the goal of creating a reusable World Wide Web-based catalogue framework for the
management of mark–recapture data accumulated by the global research community,
ecotourists, government agencies and citizen scientists. The Shepherd Project effort was
completed in 2004 and first employed in the ECOCEAN Whale Shark Photo-identification
Library (see www.whaleshark.org). The library is a repository for whale shark spot-pattern
16
data and the photographs from which they are derived. Basic information required to
accompany photographs includes: (i) sighting date and location, (ii) sex and size of the
animal, and (iii) contact details of the submitter. The library also employs two pattern-
matching algorithms (modified Groth and I3S) (Holmberg et al. 2009).
This library serves as a visual database of whale shark (R. typus) encounters and of
individually catalogued whale sharks (Arzoumanian et al. 2005). The library is maintained
and used by researchers to collect and analyse whale shark encounter data in order to explore
various aspects of the species and its life history. The library uses photographs of the skin
patterning behind the gills of each shark and any scars to distinguish between individual
animals (Appendix 1). Cutting-edge software supports rapid identification using pattern
recognition and photo management tools. Importantly, it is available for use by researchers
working on whale sharks throughout the world (see Appendix 1).
Because pattern-recognition is a resource intensive task, the ECOCEAN Library
utilises a ‘sharkGrid’ to distribute tasks between multiple computers to allow scans to be
completed in a timely manner. The library further utilised the support of Murdoch University
and the Pawsey Supercomputing Centre (an unincorporated joint venture of CSIRO and the
four public WA universities with funding from the Western Australian Government) to join
sharkGrid and process the pattern- recognition scans for identifying whale sharks. More
recently, this role has been allocated to Amazon Elastic Compute Cloud (Amazon EC2),
which is a web service that provides resizable compute capacity in the cloud.
Initiated from whale shark sightings in Western Australia (Norman 1999), the earliest
entry to the ECOCEAN Library was from underwater film maker Ben Cropp who
photographed a whale shark off Montague Island, NSW in 1964. This individual was
allocated the codename A-600, one of more than 1100 individuals identified from Australian
waters. To further encourage community uptake, especially at locations throughout the
17
world, ECOCEAN initiated an outreach program (with support from the Rolex Award for
Enterprise program) where information posters (see Figure 1.1) were distributed to more than
100 dive shops in Australia and a further 20 countries.
Figure 1.1: Whale shark photo-identification program posters.
The whale shark photo-identification library has expanded considerably since its inception
(Figure 1.2), and currently stores more than 50000 whale shark identification images. By
2013, more than 28000 individual whale shark sighting reports are currently housed in the
database (also known more recently as Wildbook for Whale Sharks) with data on more than
6000 individual whale sharks submitted from contributors in 54 countries.
18
Total Whale Shark sighting reports
Year
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
Num
ber o
f sig
htin
gs re
porte
d
0
5000
10000
15000
20000
25000
30000
155 6261509
2558
4354
6801
10052
17677
14312
22044
25788
Figure 1.2: Growth of the ECOCEAN Whale Shark Photo-identification Library globally
(2003-2013).
Hypotheses and aims of the study
The unique, individual spot patterning found on each whale shark, combined with their
longevity, will allow a number of hypotheses to be tested during this study, utilising citizen
science together with photo-identification and contemporary tagging techniques. Firstly,
these techniques will allow the hypothesis that the species is wide ranging and undertakes
large inter-continental migrations to be tested. Secondly, some aggregations, such as that in
Western Australia, will be seasonal and comprised of individuals that return each year during
periods of high prey concentrations, then undertaking large-scale migrations during other
periods of the year. Photo-identification and ‘citizen science’ can be used to effectively
monitor whale sharks on a global scale and to determine the number of individuals at specific
aggregation sites. The addition of contemporary electronic tagging techniques will allow the
19
determination of short term movement patterns within and beyond aggregation sites to
determine home range and residency patterns in Western Australia. It is therefore the aim of
this thesis to employ a combination of telemetric studies, contributions from citizen scientists
and the unique patterning of whale sharks to examine global aggregations and movements
between specific whale shark ‘hotspots’, as well as determining the movement patterns and
population characteristics of the species within Western Australian waters.
20
Chapter 2
Undersea constellations: ‘Citizen Scientists’ elucidate the global
biology of a threatened marine mega-vertebrate
Introduction
Gathering data on the life of enigmatic animals, particularly on behaviours, habits and
movements, remains a challenge, despite its important role in biodiversity conservation and
management. For many species, biogeographic investigations are largely the result of
information that is generated from multiple sources, often over long time-scales, because
measuring biogeographic and biological data over large geographic areas is simply not
feasible by a single team of researchers (Chiarucci et al. 2011). One approach that has proven
promising in addressing many of these issues is ‘citizen science’ (Bonney et al. 2009,
Newman et al. 2012). In the age of a well-educated public and accessible and mobile digital
technology, scientists are able to harness the observations of millions of people, thus greatly
increasing their power of observation (Newman et al. 2012). For many charismatic species,
public awareness is high but numbers of study species can often be low, particularly for
threatened species, and citizen science has the potential to provide a powerful tool for
biological investigation. The current study explores how citizen science can contribute to our
understanding of the basic biology and ecology of the whale shark (Rhincodon typus) on a
global scale.
Relatively few sightings of whale sharks were recorded in the literature prior to the
mid-1980’s (Wolfson 1986). Indeed, many of the whale shark aggregation sites have only
21
been documented in the past decade (Rowat and Brooks 2012, Pierce and Norman 2016).
Whale sharks are one of only three filter-feeding species (Motta et al. 2010). They are known
to aggregate, generally in groups (or constellations) of juvenile males at hotspots/regions
throughout the world’s oceans where their planktonic prey may seasonally mass (Compagno
1984, Colman 1997, Riley et al. 2010, de la Parra et al. 2011, Rohner et al. 2013, Vignaud et
al. 2014). Whale sharks are distributed throughout the world’s oceans in the region between
30oN and 30oS latitude (Last and Stevens 1994), and exhibit "K" selected life history
characteristics, which includes slow growth, late maturation and extended longevity (Colman
1997). These are a few of the traits responsible for their listing as ‘Vulnerable (VU)’ under
the World Conservation Union (IUCN) Red List of Threatened Species (Norman 2000).
In recent years, improved monitoring techniques and the upsurge in ecotourism
industries centered on this species, have ensured that biological and ecological information
has increased significantly (Arzoumanian et al. 2005, Stevens 2007), enabling an improved
understanding of the primary locations and the timing of whale shark appearances throughout
its range. Satellite telemetry and bycatch data are now informing which environmental factors
may drive the formation and dissolution of such aggregations (Wilson et al. 2001a, Sleeman
et al. 2010, Sequeira et al. 2012). However, the sample sizes of whale shark tagging studies
remains low (see e.g. Eckert and Stewart 2001, Wilson et al. 2006, Graham et al. 2006,
Gifford et al. 2007, Sleeman et al. 2010, Hearn et al. 2013).
The use of photo-identification in whale shark monitoring provides an opportunity to
‘tag’ an animal in a non-invasive manner and ensure this ‘natural tag’ is available for use in
long term resighting programs (Arzoumanian et al. 2005, Graham and Roberts 2007, Speed et
al. 2007, Rowat et al. 2009, Marshall and Pierce 2012). The photo-identification system
utilises the natural skin patterning on whale sharks to identify individual animals (Taylor
1994, Norman 1999). A database of photo-identified whale sharks was created in 1995 from
22
data collected at Ningaloo Marine Park, Western Australia (Norman 1999). The Wildbook
for Whale Sharks (founded as the ECOCEAN Whale Shark Photo-identification Library
(www.whaleshark.org)) was published online in 2003 to enable easy submission of whale
shark sighting data from ecotourists (citizen scientists) and researchers. This portal serves as
a globally and regionally scoped research platform for standardised capture-mark-recapture
studies (Holmberg et al. 2008, 2009) and provides a unique opportunity for global
collaborations across contributing scientists.
Here, the success of the global monitoring of whale sharks is reported and the
potential of the Wildbook database is explored, in both capturing global aspects of whale
shark biology, including regionally explicit population characteristics, such as sex-ratios and
size compositions. The efficacy of large scale citizen science efforts to provide key
information regarding the life-history of a charismatic species is highlighted, with a
discussion following on the potential biases and challenges in the implementation of a
research program involving the general public.
Search functions within the library were employed to: i) undertake an extensive
review of whale shark sightings over an extended period at the local and global level; ii)
determine resightings of individual whale sharks in one or more countries; iii) establish the
top locations with extended resight history for 20 or more individual whale sharks; and iv)
establish size and sex ratios of these locations over an extended period.
23
Materials and Methods
The primary database with information on whale shark sightings worldwide used in this study
was the ECOCEAN Whale Shark Photo-identification Library/Wildbook for Whale Sharks
(www.whaleshark.org, Arzoumanian et al. 2005). Whale shark identification images are
collected when a swimmer photographs the individual’s unique spot pattern immediately
behind the gill slits (see Appendix 1, Figure 1.1, Arzoumanian et al. 2005), which is distinct
and long-lasting (Marshall and Pierce 2012), and this image is then submitted to the online
library. Participants also upload, where possible, other relevant sighting information to the
database for storage and future analysis such as the sex and estimated total length (TL).
While length estimates can vary depending on experience of the recorder (see Rohner et al.
2011), repeat sightings of identified individuals provides increased confidence that the correct
sex for each animal has been accurately determined.
Researchers working in the various aggregation sites process the appropriate images
as described in Arzoumanian et al. (2005). Computer-assisted scanning technology is used to
determine whether the individual whale shark in question is a ‘new’ shark or a ‘resight’ of a
previously reported whale shark within the database. Each encounter is automatically
assigned a location code, depending on the country or region where the encounter occurred.
An ‘encounter’ is defined as a whale shark sighting with information on the location,
preferably combined with an associated identification photograph that has been submitted to
the library. These data are then shared between all interested parties via the global online
database, enabling international matches (and therefore movement between locations) to be
determined. As not every whale shark encounter submission has an identification photograph
of suitable quality to confirm the individual shark’s identity, some encounters remain
unassigned to a specific shark identity. Identified sharks are catalogued with a prefix
24
according to the locality of first identifiable sighting (e.g., ‘A’ for Australia, ‘BZ’ for Belize)
and each newly identified shark is assigned a unique number specific to that sighting location
(e.g., A-001, A-002, BZ-050, BZ-051 etc.).
Results
Global hotspots/regions for whale sharks
From 1992 to 2014, the library had received a total of 28776 whale shark encounter reports
resulting in the identification of 6091 individual whale sharks from 54 different countries.
For this study, the primary datasets used were from the top 20 whale shark hotspots/regions
with >100 encounters recorded in the library for the period spanning 1992-2014 (see Figure
2.1), where 28529 encounters (99.14%) were received resulting in the photo-identification of
5955 (or 97.77%) of all individuals (Table 2.1). Thus the number of whale shark encounters
submitted from across the globe continued to increase from the moment the library was
published online in 2003, although some sightings that predated it were also available for
inclusion in the dataset (Figure 2.2).
Uptake of the library was not uniform at all global hotspots/regions, with Ningaloo
Reef, USA Gulf States and Thailand representing the locations with the earliest data
submissions (1992) and more recently from Tanzania (2006). However, the level of uptake at
each hotspot/region has generally been more intensive in recent years (Table 2.3, Figure 2.2).
The locations with the greatest number of unique individuals identified via photo-
identification were Mexico (Atlantic) (n=1101), Ningaloo (Western Australia) (n=1082),
Philippines (n=775) and Mozambique (n=676) (Figure 2.3).
25
Sex ratio
Based on the submission of images to the photo-identification library, there is a
predominantly male bias at the large majority of sites, with few exceptions. At the Galapagos,
99% of sexed individuals were female, while in the Red Sea, 75% were female, and in
Thailand, 68.5% were female (Figure 2.4). This contrasts markedly with a number of
locations, for example in the Maldives and South Africa where only 9.43% and 9.60 %,
respectively, of the sexed whale sharks that were submitted to the photo-identification library
were females (Figure 2.4). However, at 14 of the top 20 global whale shark aggregation sites
>66% of the identified whale sharks were male.
Size
Mean total length (TL) at the different locations varies, with the largest occurring at the
Galapagos (mean = 11.07 m TL (± 0.30 SE)), followed by the USA Gulf States (mean = 8.01
m (± 0.28 SE)), Belize (mean = 7.21 m (± 0.24 SE)), and Mexico (Atlantic) (mean = 7.12 m
(± 0.06 SE)). All other locations reported a mean TL that was less than 7.0 m, the smallest
whale sharks being found in Thailand, Djibouti and Indonesia, where TL <4.6 m (Table 2.2).
The size (TL) of maturity of whale sharks in the Indo-Pacific population has been determined
to be around 8.1 m in males (Norman and Stevens 2007), while the Atlantic population may
be mature at somewhat smaller sizes for both males and females (see Hueter et al., 2013).
Site fidelity
Across the 20 global hotspots/regions, whale sharks are found in relatively high numbers at
some localities throughout most of the year (e.g. Maldives, Mozambique, Thailand, Red Sea,
26
Honduras) (Figure 2.6). Shark M-014 was recorded in the Maldives in January, February,
April, May, June, August, November and December 2008; while M-070 was recorded there
in April, August and December 2014, and M-084 was recorded in January, February, April,
August and November 2014. This suggests individual whale sharks may remain in the
Maldives throughout the year. At most hotspots/regions, constellations are highly seasonal,
e.g. Ningaloo Reef (Western Australia), Mexico (Atlantic), Belize, Philippines, Seychelles,
Tanzania and Christmas Island where sightings are restricted to less than six months of the
year (Figure 2.6).
Within each of the 20 global hotspots/regions, the percentage of identified sharks that
were observed in two or more years was calculated (Table 2.1). At Belize, the greatest
percentage of returning individuals (76.6% of the 47 individual sharks identified) was
recorded, followed by Maldives (60.4% of 101 sharks) and South Africa (60.0% of 45
sharks), whereas whale sharks from the Galapagos Islands showed the least evidence for site
fidelity, only one of 141 identified sharks resighted in any year subsequent to initial
identification (Table 2.1). For the 20 hotspots/regions analysed, the overall mean percentage
of sharks returning in two or more years is 35.7%.
Although the number of years the database has been populated differs among sites
(see Figure 2.2), it has been possible to establish that long-term site fidelity is present at
Ningaloo Reef with one shark (A-103) resighted over a 21 year period. Other locations with
extended site fidelity include Belize (15 years), Honduras (12 years), Mexico (Atlantic), the
Philippines, and the Seychelles (11 years), while the least maximum number of years
between resightings is in the Galapagos and Christmas Island (1 year) (Table 2.3).
27
International resightings
Photo-identification has demonstrated the movement of a number of individual whale sharks
between countries (Appendix 2), although of note was A-424 (recorded as having moved the
greatest minimum one-way distance i.e. 2700 km between Australia and Indonesia over a 4
year period) and H-021 (recorded at 4 different countries spanning 1300 km (i.e. Belize,
Honduras, Mexico (Atlantic), and USA) over a 14 year period). In addition, 6 individuals
were recorded moving between Belize, Honduras and Mexico (Atlantic) (1 over 14 years); 13
individuals moved between Belize and Honduras (1 over 13 years); 9 moved between Belize
and Mexico (Atlantic) (1 over 12 years); shark BZ-010 was resighted moving between
Belize, Mexico (Atlantic) and USA over a 10 year period; 9 sharks moved between USA-
Mexico (Atlantic) (1 over 8 years). Sharks were also recorded moving between USA-
Honduras, South Africa-Mozambique, Mozambique-Tanzania, Seychelles-Tanzania, Saudi
Arabia-Djibouti, Mexico (Atlantic)-Cuba, Oman-Qatar, Oman-United Arab Emirates, and
Taiwan-Philippines (Appendix 2).
Discussion
Global hotspots
Whale shark ecotourism has expanded globally since it was first pioneered in Western
Australia (Colman 1997). With this expansion has come an increase in whale shark sightings
(DPaW 2013). An easily accessible global database to store whale shark sightings was not
available until 2003 when the ECOCEAN Library became the central database employed for
this purpose. The extent to which the library was populated for each location however was
staggered dependent on community education. This enabled an expansion of outreach and
28
training efforts focusing on many whale shark aggregating sites and subsequent acceptance
by researchers and/or managers (Figure 2.2) and ensured a robust dataset was available for a
review on the biology and ecology of this species.
The relatively recent expansion of citizen science monitoring of whale shark
populations around the world has enabled a significant expansion in the number of
recognised global hotspots/regions for this species to be increased from 13 to 20 (e.g. Rowat
and Brooks 2012, Berumen et al. 2014) (see Figure 2.1). However, three of the four countries
with historically the most extensive targeted fishery for this species (i.e. Taiwan, India and
China) (Pierce and Norman 2016) have not been included in this list as data from photo
monitoring studies for each is limited. While whale sharks are protected in each country, a
targeted fishery still exists in China (Li et al. 2012), with anecdotal reports of illegal catches
in several other countries. The uptake of a dedicated monitoring program is required to
establish the population demographics of whale sharks at these locations.
Despite a sex ratio at birth of almost 1:1 (Joung et al. 1996), aggregations of whale
sharks at coastal hotspots/regions (Figure 2.1) appear dominated by immature individuals of a
small to medium size (Figure 2.5) and have a generally male bias (Figure 2.4) (Norman and
Stevens 2007, Graham and Roberts 2007, Araujo et al. 2014). Exceptions can be found at
smaller aggregation sites, such as the Saudi Arabian coast of the Red Sea where there is a
non-biased (1:1) male to female ratio (Berumen et al. 2014), and at the Azores where large
(>8 m TL) individuals dominate (Afonso et al. 2014), and offshore at the southern Gulf of
California and at the Galapagos Islands where large, possibly pregnant, females are common
(Ramirez-Macias et al. 2007, Acuna-Marrero et al. 2014). Sex and size segregation is not
uncommon amongst shark populations (Klimley 1987, Ketchum et al. 2013, Vandeperre et
al. 2014) and it has been documented in >10% of species for which biological data are
available (Compagno 1984); and has been related to sex differences in body size,
29
reproductive cycle, predation risk, forage selection, activity budget, behaviour, thermal-niche
fecundity and social factors (Wearmouth and Sims 2008, Kock et al. 2013). Interestingly,
records of whale shark neonates are limited and pupping and nursery areas remain
unidentified (Rowat and Brooks 2012). It has to be noted, however, that some species of
shark do not use geographically restricted nurseries and pupping may occur over large
geographic areas (Heupel et al. 2007), especially for whale sharks given the way the young
appear to develop (see Schmidt et al. 2010).
Peak sighting periods and site fidelity
Sighting peaks within the current study often occurred around the same time as peaks in
plankton abundance, although search effort, being closely tied to ecotourism activities, tended
to focus around these times in order to maximize success. . Whale shark aggregations often
coincide with productivity events (Graham et al. 2006, Sleeman et al. 2010, de la Parra
Venegas et al. 2011, Ramirez-Macias et al. 2012), which can be high for either a short or
long period, thus providing significant feeding opportunities (Nelson and Eckert 2007) that
are often exploited by whale sharks on an annual basis (Taylor 1994, Colman 1997, Duffy
2002, Graham et al. 2006, Hoffmayer et al. 2007, Stevens 2007, Taylor 2007, de la Parra
Venegas et al. 2011, Fox et al. 2013, Gleiss et al. 2013, Robinson et al. 2013).
During feeding, total energy requirements can be met in a few or several hours (Motta
et al. 2010), with Gleiss et al. (2011b) suggesting that even short periods of active feeding (8
min/day) on exceptionally high concentrations of prey may satisfy the energy requirements of
whale sharks. This behaviour relative to production and prey availability has previously been
hypothesized as the reason for distributional shifts for both basking sharks (Sims and Reid
2002) and whale sharks (Graham 2007, Rohner et al. 2013).
30
There is evidence for long-term site fidelity among whale sharks at multiple global
hotspots/regions (up to 21 years at Ningaloo Reef, Western Australia, for example) is
occurring, with many identified whale sharks within these feeding aggregations returning to
the same location in subsequent years (Table 2.1). Barendse et al. (2011) reports that in a
photo-identification study of humpback whales, a resighting rate of 15.6% at intervals of one
or more years indicates long-term fidelity to a particular region. Accordingly, for the top 20
global hotspots, the fact that approximately one third of all whale sharks return to a familiar
site in a subsequent year(s) indicates strong site fidelity in this species. Whale sharks appear
to have the ability to prepare for and target prey aggregations (Gunn et al. 1999, Graham et
al. 2006, Gleiss et al. 2013, Schleimer et al. 2015).
In Mozambique, Maldives and Honduras, there is clear evidence of year-round whale
shark presence (see Figure 2.6). However, despite the ecotourism industry undertaking whale
shark tours throughout most months of the year in Mozambique, none of the >600 identified
whale sharks were resighted over a period in excess of six months in any one year (although
MZ-169 was resighted on two days separated by a 4.5 month period). In contrast, in the
Maldives, citizen-science based photo-identification within this study has been used to
confirm at least some sharks have a year-round residency.
Animals move to fulfil their basic biological goals of gaining energy, seeking safety,
learning, and reproducing (Nathan et al. 2008). In the case of whale sharks, the
predominance of small and immature individuals evident at most aggregations studied
(Figure 2.3) appears to coincide with important regular natural feeding opportunities,
although the prey items are somewhat varied between constellation sites close to the relative
safety of a coastal environment (Clark and Nelson 1997, Norman 1999, Heyman et al. 2001,
Jarman and Wilson 2004, Hoffmayer et al. 2007, Nelson and Eckert 2007, Meekan et al.
2009, de la Parra Venagas et al. 2013, Fox et al. 2013, Gleiss et al. 2013, Robinson et al.
31
2013, Rohner et al. 2013). While individual whale sharks are small and immature, the prime
directive for members of these aggregations may be to expend minimal effort to find food
and increase in size and relative fitness (especially to avoid predation) prior to expending
greater energy reserves in the search for mates and reproduction. This may be achieved by
exploiting shallower coastal aggregations of prey. Exactly where the individuals reside for
the rest of the year remains largely undefined, although it is possible that whale sharks are
present but simply unavailable for capture by photo-identification monitoring techniques
(Cagua et al. 2015). In addition, it is possible that larger individuals may have an increased
ability to forage deeper into the epipelagic and mesopelagic zones (Thums et al. 2012,
Wilson et al. 2006). Although in India for example, Borrell et al. (2011) used stable isotope
profiles to suggest that sharks smaller than 4 m TL feed in a pelagic offshore habitat prior to
coming to inshore areas as they grow, while in the Gulf of California small juveniles
aggregate to feed in coastal waters of the bays and adult females feed offshore (Ramirez-
Macias et al. 2012a). Rohner (2012) suggests that whale sharks in Mozambique prey on
demersal plankton, deep sea crustaceans and fish, in addition to surface coastal zooplankton.
International resightings
Despite the apparent level of site fidelity evident in this study, a limited number of
individuals have been confirmed moving between one or more nearby countries via: marker
tags e.g. Seychelles/Mozambique (Rowat and Gore 2007); photo-identification e.g.
Belize/Mexico(Atlantic)/Honduras/USA (Hueter et al. 2013); and satellite tracking studies
e.g. Cuba/Mexico/Belize/Honduras (Graham et al. 2007); Taiwan/Japan/Philippines (Hua
Hsun Hsu, Department of Environmental Biology and Fisheries Science, National Taiwan
Ocean University, Keelung, Taiwan, personal communication);
Madagascar/Mozambique/Seychelles (Rachel Graham, MarAlliance, personal
32
communication); Australia/Indonesia (Sleeman et al. 2010); Mozambique/Madagascar
(Brunnschweiler et al. 2009); Utila/Belize/Mexico(Atlantic) (Gifford et al. 2007);
Mexico/Saint Peter and Saint Paul Archipelago, Atlantic Ocean, Mexico(Atlantic)/Cuba
(Hueter et al. 2013); and Saudi Arabia/Egypt/Yemen/Oman (Berumen et al. 2014). On
occasion, these movements can occur over a very short timeframe: H-001 was photographed
in Honduras in 2005 and in Belize three days later; MZ-494 was sighted in Mozambique in
2011 and resighted within 16 days in South Africa; while BZ-026 was photographed in
Mexico (Atlantic) and Honduras within a period spanning three months. However, most of
these movements are relatively small (<1000 km) and although it is commonly accepted that
whale sharks are highly migratory (IUCN SSG/CMS 2007), few reliable records exist for
extensive movements across ocean basins (Hueter et al. 2013).
Long-distance migration of individuals within some species to exploit favourable
feeding opportunities is, however, well documented, including birds (Elphick 2007), turtles
(e.g. Chelonia mydas) (Luschi et al. 1998) and whales (e.g. Orcinus orca) (Pitman and Ensor
2003). The current study has confirmed that at least some individuals within whale shark
aggregations undertake longitudinal movements, albeit at the largely sub-adult life stage, and
usually at coastal margins. Given favourable prey availability at each location (Sleeman et
al. 2010, Rowat and Brooks 2012), these movements are potentially driven by feeding
opportunities.
Sequeira et al. (2013) summarised a limited number of published reports to suggest
that whale shark appearance timings at locations in the Indian Ocean appear sequentially,
proposing a broad movement of individuals from South Africa to Ningaloo, Western
Australia. However, despite more than 6000 individual whale sharks identified at coastal
hotspots/regions worldwide from data supplied from >4000 individual researchers and citizen
scientists and collated within the ECOCEAN (Wildbook) database, there remains no matched
33
sharks between these different continents. It therefore seems unlikely that the concept of a
broad movement of coastal (young and immature) whale sharks occurs. Rather, it is likely
that prior to the onset of maturity, whale sharks take advantage of coastal feeding
opportunities, and then as they mature, at least some may engage in more extensive
migrations from each population while generally remaining within their native ocean basin as
suggested within a recent genetic study (Vignaud et al. 2014). Genetic studies to date have
indicated that some level of trans-ocean mixing does occur between animals found within the
Pacific and Indian Oceans, while this mixing is at reduced levels between Indian/Pacific and
Atlantic Ocean animals (Jennifer Schmidt, University of Illinois, personal communication).
Because of the paucity of large, mature individuals present at these coastal aggregations
however, opportunities to investigate such movements via photo-identification or satellite
tracking are extremely limited. Nonetheless, the present study using photo-identification
does demonstrate linked connectivity among a number of coastal aggregation sites.
According to Heupel et al. (2007), shark nursery areas are defined as having i) a
greater abundance of young of the year sharks than other areas; ii) individuals displaying a
tendency to remain or return for extended periods; and iii) individuals using the area
repeatedly across years. Since most hotspots/regions identified within the current study
exhibit criteria ii) and iii), these can subsequently be defined as important ‘post-nursery
conditioning areas’. Given the high proportion of immature male animals (<8 m) within
coastal aggregations (e.g. Graham and Roberts 2007, Norman and Stevens 2007, Rowat et al.
2008, Bruunschweiler et al. 2009, Rohner et al. 2013, Fox et al. 2013, Hueter et al. 2013,
Rohner et al. 2015) the ultimate ‘need to feed’ to attain a large size is possibly the main
driver for whale sharks to aggregate and return to exploit known feeding opportunities at
these locations.
The reproductive biology and mating habits of whale sharks remains elusive, with few
34
clues based on chance encounters. Neonate records from the Philippines (Aca and Schmidt
2011), Taiwan (Hsu et al. 2014), the northern Indian Ocean (Rowat et al. 2008), St Lucia in
the Caribbean (https://www.facebook.com/SCUBASTLUCIA/?fref=photo), and the Maldives
(http://www.haveeru.com.mv/news/57774) combined with the capture of the pregnant
individual off Taiwan (Joung et al. 1996) may indicate a pupping area close to these
locations. However, staggered (see Schmidt et al. 2010) and potentially long gestation
strongly argues against specific pupping grounds, as does the fact that any neonates found
have been singles and perhaps doubles at most. Hueter et al. (2013), Ramirez-Macias et al.
(2007, 2012a), Ketchum et al. (2013) and Hsu et al. (2014) have suggested that offshore
habitats may provide pupping and nursery areas for whale sharks. Large females are presently
found in the Southern Gulf of California, the Galapagos, and St. Helena islands. Interestingly
however, only one possibly mature female from the Southern Gulf of California has been
recorded revisiting that location after seven years (Ramirez-Macias unpublished data). Only
one individual has been recorded revisiting the Galapagos in subsequent years. Long-term
monitoring may shed further light and help solve some of these mysteries.
The onset of maturity and the concomitant urge to find a suitable mate may be the
catalyst to drive larger scale movements of individual whale sharks from predominantly sex
and size segregated coastal resident aggregations where known feeding opportunities exist. It
is at times and locations when whale sharks aggregate (especially at coastal constellations)
that they may become susceptible to illegal, unregulated and unreported (IUU) fishing
pressure, which may rapidly become unsustainable for the species unless addressed (Norman
2000). In addition, shark species have discrete locations for pupping, nursing and mating
(Vandeperre et al. 2014) and identification of these essential habitats is imperative when
designing appropriate management regimes (Gruss et al. 2011). For whale sharks, this
demands greater attention and continued collaborative efforts by international stakeholders to
35
define migration routes, timings of movements, and especially critical breeding and pupping
locations.
The current study has highlighted the benefit of engaging citizen scientists, eco-tour
operators and specific researchers in the use of photo-identification to monitor whale sharks
on an international scale. This non-invasive technique is long-lasting and will enable the use
of mark-recapture analysis to monitor trends in sighting numbers initially at specific
constellation sites. These results will then be available to analyse collectively to underpin the
development of a global assessment of whale sharks throughout the species range.
While having numerous benefits, the technique is however dependent on the
collection of suitable images for use with the photo-recognition software and it also requires
adequate sampling to ‘capture’ sightings outside popular tourism periods. As such, there may
be some areas frequented by whale sharks that are yet to be adequately sampled. To address
this data gap, directed research programs should dedicate their efforts to photo-identification
sampling in areas outside of popular tourist destinations. Importantly, all monitoring data
will remain securely stored within Wildbook and available to assist the development of future
national and international management plans aimed at ensuring the long-term conservation of
the whale shark.
36
Figure 2.1: Global whale shark hotspot/regions (1-Ningaloo Marine Park; 2-Mexico Atlantic;
3-Mozambique; 4-Philippines; 5-Seychelles; 6-Honduras; 7-USA-Gulf States; 8-Maldives; 9-
Mexico Pacific; 10-Thailand; 11-Djibouti; 12-Galapagos; 13-Belize; 14-South Africa; 15-
Tanzania; 16-Oman; 17-Qatar; 18-Red Sea; 19-Christmas Island; 20-Indonesia). Coloured
groupings represent hotspots/regions within which international whale shark movements
have been confirmed via photo-identification (i.e., between 2, 6, 7, 13; between 3, 14, 15;
between 5, 15; between 16, 17; between 11, 18; and between 1, 20). N.B. One identified
whale shark has also been recorded at both Taiwan and Philippines; and another at both
Thailand and Malaysia.
37
Figure 2.2: The cumulative number of encounters submitted into the whale shark photo-identification library by the top 20 sighting locations
(from www.whaleshark.org).
38
Global Hotspot
Ningalo
o
Mexico
Atlanti
c
Mozam
bique
Philipp
ines
Seych
elles
Hondu
ras
USA - Gulf
Stat
es
Maldive
s
Mexico
Pac
ific
Thaila
nd
Djibou
ti
Galapa
gosBeli
ze
South
Africa
Tanza
niaOman
Qatar
Red S
ea
Christm
as Is
land
Indon
esia
Num
ber o
f ide
ntifi
ed s
hark
s
0
200
400
600
800
1000
1200
Figure 2.3: Total number of individual whale sharks identified in each global hotspot/region
(1992-2014).
39
Global Hotspot
Ningalo
o
Mexico A
tlanti
c
Mozambique
Philippine
s
Seyche
lles
Hondura
s
USA - Gulf
Stat
es
Maldive
s
Mexico P
acific
Thaila
nd
Djibouti
Galapag
osBelize
South A
frica
Tanza
niaOman
Qatar
Red Sea
Christm
as Is
land
Indone
sia
Perc
enta
ge
0
20
40
60
80
100
FemalesMales
529 344 492 460 163 80 45 53 236 54 41 118 22 31 115 8 277 20 16 19
Figure 2.4: Sex ratio for identified whale sharks at global hotspot/regions (1992-2014).
40
Global hotspot
Ningalo
o
Mexico
(Atla
ntic)
Mozam
bique
Philipp
ines
Seych
elles
Hondu
ras
USA (Gulf
state
s)
Maldive
s
Mexico
(Pac
ific)
Thail
and
Djibou
ti
Galapa
gosBeli
ze
South
Africa
Tanz
aniaOman
Qatar
Red S
ea
Christm
as Is
land
Indon
esia
Tota
l len
gth
(m)
0
1
2
3
4
5
6
7
8
9
10
11
12
13FemalesMales
Figure 2.5: Mean TL of male and female Rhincodon typus identified within the whale shark
photo-identification library at 20 global hotspot/regions (1992-2014).
41
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Belize
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Tanzania
Oman
Qatar
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Christmas Island
Indonesia
Figure 2.6: The combined weekly patterns of whale shark encounters recorded at global
hotspot/regions.
42
Table 2.1: Site fidelity at global hotspot/regions (1992-2014).
Global Hotspot/region Total number of sighting reports
(encounters)
Total number of sharks identified
Total number of sharks sighted in 2 or more calendar years
% of identified sharks sighted in 2 or more
calendar years Belize 256 47 36 76.6 Maldives 747 101 61 60.4 South Africa 100 45 27 60.0 Tanzania 1148 131 65 49.6 Mexico (Atlantic) 6017 1101 535 48.6 Honduras 668 136 63 46.3 Mozambique 2379 676 312 46.2 Qatar 901 341 143 41.9 Western Australia (Ningaloo Marine Park) 8586 1082 440 40.7 Philippines (Donsol, Leyte, Cebu) 3603 775 266 34.3 Seychelles 451 204 59 28.9 Djibouti 281 87 18 20.7 Oman 151 69 13 18.8 USA (Gulf States) 419 101 16 15.8 Christmas Island 131 40 4 10.0 Mexico (Pacific) 1051 567 48 8.5 Indonesia 185 71 5 7.0 Thailand 642 184 11 6.0 Red Sea 399 57 3 5.3 Galapagos 415 141 1 0.7 TOTAL 28530 5956* 2126* 35.7
*This number includes a small number of sharks that have been identified at more than one location, resulting in a final figure that is slightly greater than its aggregate total.
43
Table 2.2: Mean total length (TL, m) of whale sharks identified at each of the 20 global
hotspot/regions.
Location Mean TL SE N Indonesia 4.14 0.23 45 Djibouti 4.26 0.15 65 Thailand 4.58 0.13 118 Christmas Island 4.90 0.19 33 Red Sea 5.03 0.33 43 Ningaloo 5.28 0.06 758 Seychelles 5.49 0.09 180 Mexico (Pacific) 5.5 0.13 96 Oman 5.55 0.38 19 Tanzania 5.78 0.09 125 Maldives 5.98 0.17 91 Philippines 6.16 0.07 571 Mozambique 6.32 0.05 617 Honduras 6.48 0.15 119 South Africa 6.84 0.23 34 Qatar 6.90 0.07 297 Mexico (Atlantic) 7.12 0.06 397 Belize 7.21 0.24 35 USA-Gulf States 8.01 0.28 44 Galapagos 11.07 0.30 89
44
Table 2.3: Multi-year resights for up to 20 identified individual whale sharks at 20 global hotspot/regions (1992-2014).
Locations where sighted Period of monitoring
Number of years
monitored at this site
First year with ≥20 encounters in the library
Maximum number of
years between sightings
Shark with greatest return period
Ningaloo MP, Australia 1992-2014 23 1995 21 A-103 USA Gulf States 1992-2014 23 2009 4 GC-018 Thailand 1992-2014 23 2005 4 T-026 Seychelles 1994-2014 21 2003 11 S-028 Christmas Island 1995-2014 20 2005 1 X-001 Indonesia 1995-2014 20 2010 2 ID-068 Red Sea 1997-2014 18 2007 9 R-009 Philippines 1999-2014 16 2006 11 P-002 Maldives 1999-2014 16 2003 9 M-024, M-051 Qatar 1999-2014 16 2011 3 Q-006, Q-008 Honduras 1999-2014 16 2005 12 H-006 Galapagos 1999-2014 16 2004 1 G-009 Belize 1999-2014 16 2002 15 BZ-011 Mexico (Pacific) 2000-2014 15 2003 9 MX-188 Mexico (Atlantic) 2001-2014 14 2004 11 MXA-115 Mozambique 2002-2014 13 2005 9 MZ-013, MZ-046, MZ-197, MZ-505 Djibouti 2003-2014 12 2007 5 DJ-008, DJ-012 Oman 2004-2014 11 2009 3 OM-024, OM-043 South Africa 2005-2014 10 2008 7 SA-022 Tanzania 2006-2014 9 2008 7 TZ-001, TZ-005, TZ-009, TZ-010
45
Chapter 3
The efficacy of photo-identification for monitoring whale sharks
at Ningaloo Marine Park, Western Australia
Introduction
Photo-identification provides the means to accurately identify individual animals and
establish whether new or different individuals are being monitored (Norman 1999, Urian et
al. 2014). For species that are conservation dependant, the use of non-invasive techniques
has the potential to monitor without interrupting the natural behaviour of individuals. This
has been successfully used when studying a variety of marine animals including manta rays
(Manta alfredi), sperm whales (Physeter macrocephalus), harbour seals (Phoca vitulina),
snubfin dolphins (Orcaella heinsohni), Indo-pacific humpback dolphins (Sousa chinensis),
leopard sharks (Stegostoma fasciatum) and white sharks (Carcharodon carcharias)
(Matthews et al. 2001, Parra et al. 2006, Domeier and Nasby-Lucas 2007, Dudgeon et al.
2008, Thompson and Wheeler 2008, Marshall et al. 2011).
The unique spot patterning on the skin of whale sharks lends itself to be used for the
identification of individuals (Taylor 1994, Norman 1999). While pioneered at Ningaloo
using photographic and slide film (Norman 1999), with the broad uptake of and relative low-
cost of digital photography in recent years (Markowitz et al. 2003, Mazzoil et al. 2004) has
come the opportunity for far greater uptake of the citizen science whale shark photo-
identification monitoring program within the current study.
46
History of the current photo-identification program at Ningaloo
This research study on Rhincodon typus at Ningaloo Marine Park (NMP) began in 1995
(Norman 1999) and provided the genesis for the photo-identification program used in the
current study. In 2000, ECOCEAN formed to promote public awareness for whale shark
conservation using photo-id and helped promote participation from the public in the
ECOCEAN Whale Shark Photo-identification Library (as it became known) project. The
library was enhanced through inclusion of the Shepherd Project software in 2002 and was
hosted online at www.whaleshark.org in 2003 and uses an associated pattern matching
algorithm to confirm new or resighted individual whale sharks (Arzoumanian et al. 2005, see
Appendix 1).
The Whale Shark Ecotourism Industry (WSEI) at Ningaloo Marine Park was the first
of its type worldwide focused on this threatened species (Jones et al. 2009) providing the
opportunity for tourists to swim alongside whale sharks in their natural environment. The
industry is regulated by the Western Australian Department of Parks and Wildlife (DPaW)
(formerly known as the Department of Conservation and Land Management (CALM) and the
Department of Environment and Conservation (DEC)), and the number of licences to operate
whale shark tours has varied between 13 to 15 since 1993 (Colman 1997, Anderson et al.
2014). It continues to be regarded as an example of ‘world’s best practice’ ecotourism
(Norman 2002, Rowat and Brooks 2012) and its long-term sustainability is paramount for
stakeholders in the remote region of Western Australia (Catlin et al. 2010).
As testament to the early uptake of the whale shark photo-identification monitoring
program, Ningaloo Marine Park became the first global site to have 20 or more whale shark
encounter reports (with associated identification images) submitted to the database (see
Chapter 2, Table 2.3). With regular submissions until the present day, the library has served
47
to engage the public in a robust long-term citizen science whale shark monitoring program at
Ningaloo Marine Park and provided a means to determine whale shark return rate and
recruitment, while also enabling ongoing analysis of the health of the whale shark population
at this location.
Citizen science continues to evolve and benefit conservation research (Catlin-Groves
2012, Connors et al. 2012, Dawson et al. 2015). Implementation of a program involving the
public can provide key information to assess of the annual/seasonal timing of whale shark
appearance at Ningaloo Marine Park, allowing for the determination of previously
unidentified whale sharks at the site each year (Discovery Curve). The aim of this part of the
study was to determine the percentage of previously identified whale sharks that are returning
to Ningaloo Marine Park during the Whale Shark Ecotourism Industry (WSEI) ‘season’ each
year, by examining the number of days that each identified whale shark was available to the
WSEI between 2006 and 2013.
Materials and Methods
The Ningaloo Marine Park Whale Shark Ecotourism Industry (WSEI) undertakes operations
in waters within 5 km westward of the reef crest, using spotter planes to search for whale
sharks and direct tour boats to the area to enable ecotourists to enter the water to interact with
the sharks (Norman 1999). This provided an opportunity to join tour vessels and photograph
whale sharks for the photo-identification study, which were then processed on a daily basis to
test for a match (see Holmberg and Norman 2009). In order to educate tourists in whale shark
conservation and encourage their involvement in the whale shark photo-identification
monitoring program, >50000 whale shark public awareness brochures were produced and
distributed to ecotourists at Ningaloo between 2002 and 2013.
48
Within the Ningaloo WSEI, it is usual for each tour company to employ a
videographer/photographer to capture footage/images of the whale sharks interacted with on
any given day during the whale shark season (primarily between April–July). Although
whale shark identification images and associated sighting data at Ningaloo were available for
this study from submissions received in the whale shark photo-identification library between
1995 and 2013, with the implementation of an Earthwatch Institute volunteer-supported
program from 2006 came the opportunity to process larger datasets within the library. To
take advantage of this, a copy of the industry videographer footage from each day of the
whale shark ‘season’ for the years 2006-2009 was requested and subsequently provided by
several WSEI videographers/photographers for review. Subsequent to this, DEC/DPaW
implemented a WSEI licence condition in 2010 requiring that these data were provided by
each WSEI operator to DEC/DPaW each year and these were subsequently provided to this
study for the years 2010-2013. Video footage/identification images were subsequently
processed as per Holmberg and Norman (2009) (Appendix 1) and uploaded to the whale
shark photo-identification library (see www.whaleshark.org) with associated sighting data.
Photo-identification analysis
In order to ascertain the number of individuals identified versus search effort (number of
sighting encounters), a Discovery Curve was constructed illustrating the rate at which new
individuals were encountered. Similar analyses have been undertaken within population
studies of other marine species as an index of if and when the majority of individuals may
have been observed (Baker et al. 2006, Riley et al. 2010). To test whether there was any
significant change in the mean number of days an individual whale shark was encountered
(photographed) by the WSEI for the years 2006–2013, a one-way ANOVA was used.
49
Population estimate
Mark-recapture analysis on the whale shark population in North Ningaloo was performed
using the software program MARK (White and Burnham 1999). Capture histories were
obtained for all whale sharks identified within the whale shark photo-identification library
from North Ningaloo only between 1995 and 2013 (813 in total) as sampling effort was most
comprehensive at this site (see e.g. Holmberg et al. 2008). Detections were pooled within
each year, i.e., each shark was either detected or not detected within each year. It was
assumed as an open population as closure could not be reasonably assumed given the
migratory life-cycle and the length over which the analysis was conducted. Therefore, open
population modelling was conducted using the POPAN function (Schwarz and Arnason
1996) for the whale sharks at North Ningaloo in each year. The MARK program allowed
estimation of the population on each sampling occasion (Nj), the super-population size (N)
(i.e. total number of Whale sharks present in between the initial and final sampling occasion)
(Schwarz and Arnason 1996, 2010). The other parameters that were estimated are apparent
survival rate (fj), probability of entry to the population (bj), and probability of capture (pj).
Parameters in the models are able to be assigned as time dependant (t) or constant (.) and a
fully time dependant Cormack-Jolly-Seber (CJS) model was used initially (Cooch and White
2010).
Initially, a Goodness-of-fit (GOF) test for the model was undertaken using the
RELEASE program to validate the assumptions of the model. The latter program provides
GOF for two underlying assumptions in the CJS model, 1) Test 2 (i.e., testing the assumption
of equal capture probability between individuals) and, 2) Test 3 (i.e., testing the assumption
of equal survival probability between individuals). The results of Test 2 + Test 3 (i.e.,
rejection of their null hypotheses of adherence to their respective assumptions) may
necessitate the addition of a variance inflation factor (ĉ = χ2 (TEST 2 +TEST 3)/df) to correct
50
for the extra variation in the model that results in the estimation of quasi-Akaike Information
Criterion (QAICc).
All combinations of time-dependence (t) and consistent (.) parameters were
constructed in POPAN (i.e., eight models) and the most parsimonious chosen on the basis of
ranking their QAICc.
Results
Whale shark photo-identification library uptake
The whale shark photo-identification library at Ningaloo Marine Park (North + South
Ningaloo), since its implementation in 1995 (as measured by submissions received), has seen
a steady increase to a total of 8416 encounters until 31 December 2013 (see Figure 3.1).
Within the period 2006-2013, the suitability of identification images submitted to the library
was good, enabling on average, approximately 80% of submissions to be allocated to a shark
(either ‘new’ or ‘resight’) each year (Table 3.1).
Discovery Curve
A Discovery Curve for whale sharks sighted at Ningaloo Marine Park (North + South
Ningaloo) between 1992 and 2013 showed no asymptote (with a cumulative total of 8416
encounters identifying 1057 unique whale sharks) (Figure 3.2).
How often are individual sharks available to the WSEI?
During the period 2006-2013, the year with the greatest number of whale sharks present was
2013, with 344 individuals, while the least number identified was in 2006 (n = 84) (Table
3.2). In all years, individual whale sharks were most commonly encountered for one day
51
only (Figure 3.3), although the maximum number of sighting days for any individual in any
year was as high as 18 (for A-666 over a 101 day monitoring period in 2012) (Table 3.2).
The mean number of days that individual whale sharks were available to the Ningaloo Marine
Park WSEI between 2006 and 2013 was greatest in 2010 at 2.96 (± 0.18 SE), 2009 at 2.64 (±
0.16 SE) and 2012 at 2.42 (± 0.13 SE) (Table 3.2). A one-way ANOVA showed the mean
number of days a whale shark was encountered by the WSEI differed significantly between
years (F7, 1574=12.82, p=4.21x10-16). Tukey HSD post hoc tests revealed that 2010 was the
main driver of the ANOVA result, differing significantly to all years except 2009. Significant
differences were also found between 2009 and the four years with the lowest mean number of
days encountered per shark (2011: 1.70 (± 0.10 SE) d; 2013: 1.75 (± 0.07 SE) d; 2006: 1.76
(± 0.15 SE) d; and 2008: 1.86 (± 0.09 SE) d), and 2012 also differed significantly to 2010,
2011 and 2013 (Table 3.2).
During the period 2006-2013, the mean for the percentage of whale sharks observed,
photographed and submitted to the library on two or more days per calendar year was
46.08%, with the lowest in 2011 (35.46%) and the highest in 2010 (62.83%) (Table 3.2). For
whale sharks observed, photographed and submitted to the library on two or more days per
calendar year, the mean number of days per shark per year was lowest at 2.96 (± 0.12 SE) in
2013 and highest in 2010 at 4.12 days (± 0.25 SE) (Table 3.2).
New versus old sharks
The proportion of new versus old sharks in the library has fluctuated throughout the duration
of the monitoring period (1992-2013), with the highest percentage in the early years (1992-
1996) and the lowest in 2004, 2005 and 2011. In both 2012 and 2013, the percentage of new
sharks was approximately 50% (Figure 3.4).
52
Return rate of new sharks
Using the library, it was possible to identify the cohort of new sharks each year between 1995
and 2013 and follow their resighting rate in subsequent years (Table 3.3). On average,
resights were highest in the 1st year after initial sighting (~25%) and lowest after 18 years
(~7%), although some sharks within each cohort were consistently sighted, therefore
remaining available to the WSEI over the period (Figure 3.5).
Population estimate
Of the eight models tested, three did not converge. From the results of RELEASE, a
substantial ĉ was applied to the remaining five models (Table 3.4) due to the considerable
extra variation in the models indicating a degree of violation of key assumptions of CJS. One
other model (i.e. fj(.) p(.) b(.)) had a very large quasi deviance and was therefore excluded.
The population analysis revealed that the fj(.) p(.) b(t) model (i.e. constant apparent
survival and probability of ‘capture’ and time dependant probability of entry into the
population) was the most parsimonious based on the QAICc scores (Table 3.5). This was
more than six times better supported than the next best performing model (Table 3.5). Based
on this model, the super-population size of whale sharks in the North Ningaloo over the entire
study period was 1220 (±32 SE) (Table 3.5), showing a general population increase (Figure
3.6).
Discussion
With increased participant involvement becomes increased learning (Sin 2009) and through
the combination of public awareness via the distribution of >50000 information brochures
between 2002 and 2013, and community engagement in the whale shark citizen science
53
program (Figure 1.1), the monitoring of whale sharks at Ningaloo has developed substantially
(see Figure 3.1). The evolution and increased availability of digital camera and video
equipment has also served to encourage easier and more accurate capture and analysis of
images for use in photo-identification monitoring (Markowitz et al. 2003, Mazzoil et al.
2004, Meyer 2007a, 2007b, Lodi et al. 2009).
The quality of whale shark encounter submissions from both the public and industry
stakeholders has been consistently high as evidenced by the percentage of encounter
submissions that can be confidently allocated as a new or previously identified whale shark
(Table 3.1) using the pattern recognition software within the Photo-identification Library
(Appendix 1, www.whaleshark.org). It has therefore been successful in capturing a high
proportion of individuals present at Ningaloo Marine Park (as shown by a flattening
Discovery Curve (Figure 3.2)), and the population appears to be increasing (Figure 3.6).
A total of 813 individual whale sharks were recorded visiting North Ningaloo
between 2006 and 2013, although it is likely that a greater number is utilising the Ningaloo
Marine Park as evidenced by a lack of asymptote in the Discovery Curve (Figure 3.2) and the
overall population is much larger than all individuals identified.
Raw photo-identification and sighting data were reviewed indicating an increase in
the number of individual whale sharks identified and available to the WSEI at Ningaloo
Marine Park over the period 2006–2013. While potentially supporting the population
modelling results presented by Holmberg et al. (2008), this result may also simply be a
consequence of improved data collection resulting from increased awareness and uptake of
photo-identification monitoring techniques by stakeholders within the WSEI as the WSEI
developed.
The whale shark aggregation at Ningaloo Marine Park has been the focus of several
population studies (see e.g. Meekan et al. 2006, Bradshaw et al. 2008, Holmberg et al. 2008,
54
2009). Although the study by Bradshaw et al. (2008) reported an apparent decline in whale
shark abundance, subsequent analysis of the North Ningaloo population by Holmberg et al.
(2008) showed a slight growth in the population between 1995 and 2006. Similarly, within
the context of the current study (1995-2013), a general increase in the population at North
Ningaloo was observed (Figure 3.6). While providing annual estimates of abundance and a
superpopulation of 1220 individuals, this analysis does however include a degree of non-
adherence to the assumptions of CJS (equal survival and capture) revealed in the RELEASE
analysis (Table 3.4).
The mean number of days each shark is encountered per year within the WSEI is a
measure of the availability of whale sharks for an ecotourism interaction at Ningaloo. This
could also be used as a proxy for measuring the tolerance of whale sharks to tourism
interaction at Ningaloo Marine Park over time (Norman 2002, Sanzogni et al. 2015). In the
present study, this yearly mean fluctuated during the period 2006-2013, and although no
obvious trend was evident (Table 3.2), an expanded monitoring program combined with more
focused research on whale shark behaviour under control and ecotourism activities in
subsequent years will to assist with the long-term sustainable management of this industry,
especially as tourist numbers continue to rise (DPaW 2013).
In addition, there may be some benefit in analysing any variation in the number of
‘new recruits’ that are seen in the year subsequent to first sighting at Ningaloo Marine Park
and considering its use as a metric to test for impact of ecotourism affecting return rate of
whale sharks in subsequent years. This will be enhanced through the collection of accurate
whale shark total length data in future years and will enable a review of the size of the new
recruits entering the population annually.
Quality and comprehensive library submissions are imperative, especially as evidence
to date indicates that whale sharks are only available for ‘capture’ for very few days per
55
calendar year (Table 3.2). In order to monitor this constellation of whale sharks adequately,
the maximum opportunity for data collection should be encouraged. Photo-identification has
been identified as the most appropriate monitoring option for this species (Hodgson and
Marsh 2007, Davies et al. 2013) and the current study confirms that photo-identification is a
cost-effective, non-intrusive method for monitoring whale sharks within the Whale Shark
Ecotourism Industry at Ningaloo.
56
Year
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
Cum
ulat
ive
num
ber o
f enc
ount
ers
0
2000
4000
6000
8000
10000
Figure 3.1: Cumulative whale shark encounters from Ningaloo Marine Park submitted to the
whale shark photo-identification library from 1992 to 2013.
57
Number of 'encounters'0 1000 2000 3000 4000 5000 6000 7000 8000 9000
Num
ber o
f ind
ivid
ual s
hark
s id
entif
ied
0
200
400
600
800
1000
1200
Figure 3.2: Discovery Curve (encounters versus new sharks) for Ningaloo whale sharks
(1992-2013).
58
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
Number of days sighted
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22
Perc
enta
ge o
f sha
rks
(%)
0
20
40
60
80
100
2006 (n = 84)
2007 (n = 102)
2008 (n = 161)
2009 (n = 201)
2010 (n = 226)
2011 (n = 172)
2012 (n = 292)
2013 (n = 344)
Figure 3.3: Percentage of identified whale sharks sighted on one or more days by the Whale
Shark Ecotourism Industry at Ningaloo (2006-2013).
59
Year
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
Perc
enta
ge o
f ind
ivid
ual s
hark
s id
entif
ied
(%)
0
20
40
60
80
100
Previously identified sharksNew sharks
3 1 16 33 17 29 54 23 48 119 104 78 104 84 102 161 201 226 172 292 344
Figure 3.4: Proportion of newly identified and previously identified whale sharks in each year
at Ningaloo Reef between 1992 and 2013. (NB. Total number of individuals is listed above
each column in parenthesis).
60
Number of years present after first sighting
0 2 4 6 8 10 12 14 16 18 20
Mea
n pe
rcen
tage
of n
ew s
hark
s si
ghte
d
0
5
10
15
20
25
30
Figure 3.5: Mean (±SE) percentage of newly identified whale sharks remaining available to
the Whale Shark Ecotourism Industry at Ningaloo for each year after initial sighting (1995-
2013).
61
Year
1994
1995
1996
1997
1998
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
2009
2010
2011
2012
2013
2014
2015
Popu
latio
n es
timat
e
0
100
200
300
400
500
600
Figure 3.6: Population estimate (±SE) over time (1995-2013) of the whale shark population at
Ningaloo.
62
Table 3.1: Percentage of sighting encounters with images of a high quality and submitted to
be allocated as a new or resighted whale shark (2006-2013).
Year Total encounters Encounters suitable for identification
% Identified encounters
2006 258 215 83.33
2007 354 264 74.58
2008 589 491 83.36
2009 922 821 89.05
2010 1354 1242 91.73
2011 676 461 68.20
2012 1557 1014 65.13
2013 952 805 84.56
Mean 832.75 664.12 79.99
63
Table 3.2: Whale sharks monitored via photo-identification during Whale Shark Ecotourism
Industry encounters at Ningaloo Marine Park (2006-2013).
Year No. of
encounters per year
No. of individual
whale sharks per
year
Max. No. of days an individual
shark was sighted
% whale sharks
sighted ≥ 2days
Mean No. days per shark per year sighted ≥ 2days) (±SE)
Mean No. days per shark per
year (±SE)
2006 258 84 7 36.90 3.06 ± 0.27 1.76 ± 0.15
2007 354 102 15 43.14 3.27 ± 0.40 1.98 ± 0.21
2008 589 161 7 45.34 2.89 ± 0.13 1.86 ± 0.09
2009 922 201 16 57.21 3.86 ± 0.23 2.64 ± 0.16
2010 1354 226 17 62.83 4.12 ± 0.25 2.96 ± 0.18
2011 676 172 9 35.46 2.97 ± 0.21 1.70 ± 0.10
2012 1557 292 18 49.66 3.88 ± 0.20 2.42 ± 0.13
2013 951 344 8 38.08 2.96 ± 0.12 1.75 ± 0.07
Mean 832.63 197.75 12.13 46.08 3.51 2.18
SE ±161.56 ±31.40 ±1.69 ±3.50 ±0.08 ±0.05
64
Table 3.3: Percentage of new whale sharks that remain in the population beyond year of first sighting.
1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012
# new sharks 14 27 10 12 34 17 30 79 56 30 36 41 46 97 83 87 51 137
% resighted
1996 35.7 1997 14.3 14.8
1998 21.4 11.1 10.0 1999 35.7 29.6 10.0 41.7
2000 14.3 7.4 0.0 0.0 0.0 2001 21.4 18.5 10.0 16.7 14.7 5.9
2002 21.4 29.6 0.0 33.3 23.5 29.4 36.7 2003 14.3 22.2 10.0 16.7 17.6 29.4 20.0 24.1
2004 7.1 25.9 10.0 8.3 14.7 5.9 20.0 20.3 16.1 2005 14.3 25.9 10.0 8.3 14.7 5.9 33.3 21.5 17.9 46.7
2006 14.3 18.5 10.0 0.0 14.7 5.9 16.7 12.7 10.7 10.0 8.3 2007 14.3 18.5 10.0 8.3 20.6 0.0 13.3 11.4 7.1 33.3 13.9 14.6
2008 7.1 14.8 10.0 16.7 14.7 5.9 10.0 6.3 8.9 23.3 13.9 26.8 21.7 2009 14.3 22.2 10.0 16.7 17.6 5.9 16.7 10.1 14.3 26.7 16.7 22.0 23.9 44.3
2010 14.3 22.2 10.0 0.0 20.6 5.9 13.3 8.9 10.7 16.7 19.4 9.8 23.9 38.1 47.0 2011 14.3 7.4 10.0 0.0 5.9 0.0 10.0 8.9 12.5 26.7 13.9 14.6 19.6 32.0 27.7 16.1
2012 14.3 14.8 10.0 8.3 11.8 0.0 20.0 13.9 8.9 26.7 27.8 17.1 26.1 26.8 31.3 24.1 19.6 2013 7.1 11.1 10.0 8.3 5.9 0.0 13.3 10.1 10.7 26.7 11.1 19.5 15.2 27.8 34.9 19.5 21.6 29.2
65
Table 3.4: Goodness-of-fit results for the CJS model (from RELEASE program in MARK) of
813 Whale sharks identified in North Ningaloo between 1995 and 2013.
Test χ2 df Ĉ
2 82.27 40 -
3 203.64 31 -
3.SM 19.47 15 -
2+3 285.92 71 4.03
66
Table 3.5: Model selection criteria and parameter estimates for the five models tested in MARK for population analysis of 813 whale sharks
identified between 1995 and 2013 at North Ningaloo.
Model QAICc ∆ QAICc QAICc
weight Model
Likelihood Number of parameters
Quasi Devian
ce
Super-population
(N)
95% Confidence Intervals
Lower Upper
fj(.) p(.) b(t) 956.24 0.00 0.86 1.00 20 0.00 1220 1158 1283
fj(.) p(t) b(t) 959.83 3.60 0.14 0.17 36 0.00 1259 1177 1340
fj(t) p(.) b(t) 973.56 17.34 0.00 0.00 32 0.00 1205 1140 1269
fj(t) p(t) b(t)
980.64 24.40 0.00 0.00 49 0.00 1189 1115 1262
67
Chapter 4
Residency of whale sharks at Ningaloo Marine Park informed
using acoustic telemetry and photo-identification
Introduction
Monitoring periods of presence/absence for wildlife in the marine environment presents
challenges, especially from a logistical perspective, with fieldwork often restricted by adverse
weather conditions, the need for robust and often expensive research infrastructure, and the
ability of the target species to move potentially over great distances away from the researcher
and research site (Richardson and Poloczanska 2008). This is confounded when working on
shark species, where there is also no physiological requirement for individuals to break the
surface to facilitate data collection via physical observation, as is the case for example when
working on marine mammals (Cooke et al. 2004). Telemetry options (e.g. acoustic, satellite
and archival tagging) are being increasingly utilised, although each has its limitations, i.e.
archival tags store data but usually must be retrieved for download (Costa et al. 2012),
satellite tags need to be free of the water to transmit their signals (Jewell et al. 2011), and for
acoustic tags to succeed, transmitting tags must be within a specific range of an acoustic
receiver (Jewell et al. 2011). Citizen-based monitoring or ‘citizen science’ can provide a
non-invasive monitoring option that can be very successful in advancing scientific knowledge
(Bonney et al. 2009, Newman et al. 2012), but is dependent on direct observation.
Many species of animal have natural markings on their skin which have been proven
effective in distinguishing between individuals (see Arzoumanian et al. 2005), and can be
used to monitor a population over time (Bonney et al. 2009). A citizen-based photo-
68
identification program utilising the unique pattern of spots on the lateral surface of the whale
sharks has enabled the species to be monitored at sighting locations in Australia and abroad
(see Chapters 2 and 3, Arzoumanian et al. 2005, www.whaleshark.org). It has enabled robust
population assessment for the species (Holmberg et al. 2008, 2009) and provides an
improvement on previously employed conventional marker tags for population modeling
which have been shown to have poor retention rates (Graham and Roberts 2007, Brooks et al.
2010).
Passive acoustic technology is another option available for animal tracking and is a
sightings independent method for establishing presence or absence of tagged individuals in a
given area (Heupel and Webber 2012, Cagua et al. 2015). It involves the attachment of
external or internal acoustic ‘pingers’ to individual animals, which are then recorded by
stationary receivers deployed in a monitoring region (see Heupel et al. 2006). While
surgically implanted internal tags are more invasive and requiring capture of individuals,
these tags have a relatively long retention rate in some shark species (Papastamatiou et al.
2009, Meyer et al. 2010), while external deployments are potentially subject to abrasive
influences and biofouling which may result in premature dislodgement (Dicken et al. 2006,
Jepsen et al. 2015).
At Ningaloo Marine Park, Western Australia, the Whale Shark Ecotourism Industry
(WSEI) relies on the presence of whale sharks during the period between March and July
each year, being regarded as the annual WSEI ‘season’ when most tours are undertaken
(Anderson et al. 2014). Although possible that whale sharks are present at Ningaloo Reef
outside this period, without focussed search effort (largely restricted to industry-led
monitoring during the short WSEI ‘season’), there are limited opportunities to confirm
presence/absence. It is hypothesised that a combination of non-invasive photo-identification
and the attachment of acoustic tags can be used to monitor whale shark presence/residency at
69
Ningaloo Marine Park, and specifically, to determine the extent to which the species remains
at Ningaloo Reef beyond the WSEI season.
Materials and Methods
Acoustic telemetry
Whale sharks (n = 21) were fitted with VEMCO (VEMCO, 211 Horseshoe Lake Drive,
Halifax, Nova Scotia, Canada B3S0B9) V16 acoustic tags (Figure 4.2a), that were employed
to track the movements of tagged animals up to the expected tag life of 1237 days
(www.vemco.com). These units (V16-4H-A69-9001; External PVC attachment case), with a
nominal delay of tag transmission rates of between 70 and 150 seconds, were attached via a
15 cm wire trace to a CETA-dart head (Figure 4.2a) and deployed by handspear while
swimming alongside the whale shark. Tags were positioned either near to the 1st or the 2nd
dorsal fin (Figure 4.2b) in order to provide protection against potential dislodgement should
the whale shark rub along the ocean floor or on rocky. Each shark was photographed for
identification purposes and the image and sighting data for each individual stored within the
whale shark photo-identification library.
All whale sharks were tagged at North Ningaloo (see Figure 4.1) between 2011 and
2013 under Murdoch University Animal Ethics Permit No. W2402/11 and permits provided
by DEC/DPaW (SF009184 and No. CE003919).
Tags were removed prematurely from three (3) individuals (A-236, A-546, A-795) in
2013 by persons external to the study. Two individuals (A-546, A-579) were tagged twice
during this period (i.e. 2012 and 2013; 2011 and 2013, respectively) as tags from the earlier
deployments had been lost prior to the follow-up tagging event. Each deployment was treated
as a separate animal in the descriptive statistics.
70
The Australian Animal Tracking and Monitoring System (AATAMS) Ningaloo Reef
Ecosystem Tracking Array (NRETA) was utilised to monitor the acoustic-tagged whale
sharks. Data was sourced from the Integrated Marine Observing System (IMOS), which is
supported by the Australian Government through the National Collaborative Research
Infrastructure Strategy and the Super Science Initiative. CSIRO Marine and Atmospheric
Research (Ecosciences Precinct) were responsible for regular data download from the
acoustic receivers at Ningaloo and made this available for review within this study.
A total of 138 acoustic receivers (Vemco VR2W) are secured to the ocean floor and
arranged in four (4) main ‘lines’ predominantly perpendicular to and outside the Ningaloo
Reef; the lines are positioned as far north as Tantabiddi Passage and as far south as Coral Bay
(see Figure 4.3). Dr Andre Steckenreuter (AATAMS) advised that the distances between
receivers (between and within lines) could be calculated from GPS coordinates obtained from
the AATAMS website (https://aatams.emii.org.au/aatams/installationStation/list) and using
Google Earth (http://www.google.com/earth/download/ge/agree.html) and an online distance
calculator (http://andrew.hedges.name/experiments/haversine/) showing the ‘lines’ are spaced
at distances between 10-60 km apart, with the distance between each receiver within the
‘line’ being approximately 800 m (see Table 4.1). The NRETA was expected to detect only
whether tagged whale sharks were present at Ningaloo Reef on any given day, rather than to
provide continuous monitoring of shark movements.
Data analysis
Because no receivers within the reef detected R. typus during the study, only receivers outside
or at the edge of the reef (65 in total) were included in analyses. The total number of
detections was calculated for each acoustic array, and divided by the total number of
detections to occur within the study, in order to construct the percentages of detections that
71
occurred at each array. Because the number of receivers within an array was not consistent,
the number of detections from each array was first standardised by the number of receivers
within the each respective array. However, it needs to be noted that this standardisation may
bias results if detections/presence of R. typus are/is not evenly distributed between receivers.
As such, raw detection numbers for each array are also presented. The Mangrove Bay Array
was not directly compared to the other ‘curtain’ arrays due to differences in array
design/coverage.
A Goodness of Fit Chi-Square test was run to determine if the number of detections in
a particular acoustic array varied significantly between receivers. No test was conducted for
the 23rd parallel array due to the limited number of receivers.
Photo-identification study
For methods of photo-identification, the reader is referred to Chapters 2 and 3. A residency
index was calculated for whale sharks recorded on two or more separate days within a
‘season’, establishing the monitoring period (1st and final date of sighting) and the number of
days an individual was recorded within this monitoring period. An ANOVA was used to test
for any significant difference in the mean residency period for the years 2011, 2012 and
2013.
Results
Acoustic study
During the study period, data from a small number of receivers within NRETA were lost. Of
the receivers located outside of the reef, these primarily included those within the Mangrove
Bay Array. In addition two receivers on the Southern Line array (SL7 and 21) (see Figure
72
4.3) were down between late April through October 2011 (no detections were recorded on
these receivers throughout the study).
Residency
A total of 21 R. typus were tagged with acoustic transmitters and monitored between 2011
and 2013, with all but one of the 21 tags detected within the NRETA. During this period
2295 transmissions were detected within the study site.
The monitoring period (i.e. period between date of tag deployment and final detection
of tag) of each shark ranged between 2 to 339 days (mean = 64.7 (± 17.3 SE)). Tagged sharks
were detected on 1 to 47 (9.6 (± 2.40 SE)) individual days, which produced low residency
index values (number of days detected/number of days monitored) of between 0.09 and 0.36
(mean = 0.18 (± 0.02 SE)), when shark A-795, which was monitored for only 2 d (to reduce
bias) was removed. Tagged sharks were detected within the study site for a maximum of 1 to
7 contiguous days. However, tagged sharks went without being detected for between 4 and 86
contiguous days (mean = 26.5 (± 5.1 SE), excludes sharks detected for < 1 d) (Table 4.2).
Movement and occurrence
Tagged R. typus were detected in all months of the year (Figure 4.4). During this period the
sharks were observed to have moved frequently between arrays on the western edge of the
peninsula (i.e. between the Central Line and Northern Line arrays) (see Figure 4.3). Only
35% of the monitored sharks were detected at receivers south of the Central Line Array.
In 2011, all sharks were monitored over periods ranging from 2 to 6 months. A-411
was monitored over the longest (6 month) period (September – February) and recorded at
Ningaloo Reef in all months except November and December, with a maximum gap between
detections of 95 days.
73
In 2012, the monitoring period for each shark ranged from 1 to 11 months. Two
individuals with the longest duration (A-783 and A-815) were monitored for 6 and 11 months
respectively, with a maximum period between detections of 86 days for A-783 and 50 days
for A-815.
In 2013, regular acoustic transmissions were recorded from tagged sharks post
deployment for between 1 and 4 months, with a maximum gap between recordings for any
shark being 25 days (see Appendix 3).
Site preference
Of the arrays running perpendicular to the reef, Central Line and Northern Line recorded 39.9
(610 detections) and 41.2 % (545 detections) of all detections, respectively, after array
detections were standardised by the number of receivers within each array. Conversely, the
Southern Line and 23rd Parallel arrays only recorded 3.4 (116 detections) and 3.7 % (14
detections) of these detections. When including detections from the entire study site, the
Mangrove Bay Array, the only array to run parallel to the shore, detected 20.1% (not
standardised; 370 detections) of all detections.
Intra-array comparisons of receivers demonstrated detections between receivers to be
significantly different (p <0.05), with the greatest number of detections in an array occurring
on receivers between 1 and 3 km from the reef. The receiver with the greatest number of
detections in each array accounted for between 49.5 to 68.6% (mean = 56.7% ± 4.6 SE) of
detections within the respective array. Only in the Southern Line Array were detections more
evenly distributed between receivers, with the receiver with the greatest number of detections
only accounting for 17.9 % of the array’s detections. Although sharks were detected on most
receivers within 10 km from the reef in the Southern Line Array, no detections occurred at
the three receivers between 6 and 7.5 km from the reef (Figure 4.5).
74
Photo-identification study
Sharks sighted for two or more days at Ningaloo (via photo-identification) between 2011-
2013 were analysed, showing the monitoring period (i.e. period between date of first and
final sighting) ranged from 2-159 d in 2011; 2-138 d in 2012; and 2-221 d in 2013, with
means of 41.84 (± 5.24 SE) days in 2011; 46.94 (± 2.73 SE) days in 2012; and 55.46 (± 4.07
SE) days in 2013 (Appendix 4). The Residency Index (RI) (via photo-identification)
calculated for 2011, 2012, 2013 was 0.20 (± 0.03 SE), 0.15 (± 0.01 SE), 0.12 (0.01 ± SE)
respectively, showing a decreasing trend. A one-way ANOVA showed residency indices
differed significantly between years (p=0.0071), with TukeyHSD post-hoc test revealing that
2013 and 2011 were significantly different (p=0.0050). Although 2012 showed a lower mean
residency index than 2011, they were not found to be significantly different (p=0.1305),
similarly 2013 had a lower mean residency index than 2012, but this was also not
significantly different (p=0.2329).
Acoustic tracking and photo-identification
When comparing the number of days recorded for specific individuals within both monitoring
regimes, it is clear that acoustic monitoring ‘captures’ whale sharks for significantly more
days than is possible through photo-identification (Table 4.2, Table 4.3). A Paired t-test was
used because the data were obtained from the same individual whale sharks and showed that
the mean RI calculated through acoustic tracking (mean = 0.18 ± 0.02 SE) was significantly
higher than when calculated via photo-identification monitoring (mean = 0.075 ± 0.02 SE)
(p=0.0015).
In 2011, 50% of acoustic tagged whale sharks were recorded within NRETA on the
day of tagging; 14.3% of the newly tagged whale sharks in 2012 were recorded within
75
NRETA on the day of tagging; and 57.1% of the 2013 tagged sharks were recorded within
NRETA on the day of tagging. All sharks were ‘captured’ and confirmed as present via
photo-identification at the tagging event in each year (Appendix 3).
Discussion
Photo-identification continues to be the most comprehensive monitoring option for whale
sharks currently implemented at Ningaloo Marine Park. While it has provided data on whale
shark numbers and the confirmed resighting of identified whale sharks in successive years
(Holmberg et al. 2008, 2009), it is dependent on the WSEI operations, aerial surveillance for
spotting whale sharks, swimmers being in the water, and the sharks being at the surface to
enable sighting data and identification images to be ‘captured’ by this monitoring technique.
Monitoring is however not possible at night or during adverse sea state conditions, and
therefore whale shark numbers may be underrepresented.
Alternatively, passive acoustic tracking is not reliant on direct observation (e.g. via
photo-identification) to assess presence/absence and therefore provides an opportunity to
determine the period whale sharks remain at Ningaloo Marine Park. The main acoustic
receiver lines within NRETA are spaced at distances of between 10 and 60 km apart, with the
distance between each receiver within the ‘line’ perpendicular to the reef approximately 800
m. Given the ‘range’ of the acoustic tags relative to receiver is usually in the range between
100–500 m (Heupel and Webber 2012, Cagua et al. 2015) and acoustic signal can be
influenced by abiotic factors such as wind and waves (Heupel et al. 2006), there is the
potential for a shark to be in the area but not ‘captured’ by the receiver array, thereby giving
an impression of non-presence even though the tagged sharks are in the general area.
In an earlier study, active acoustic tracking of a whale shark at Ningaloo Marine Park
76
revealed the short-term (i.e. <24 hour) movements by this individual was essentially along-
shore and parallel to the reef front (Gunn et al. 1999). The shark (A-020) was tracked during
two separate periods over a 40 km stretch and no further than 3 km westward of Ningaloo
Reef. Given Anderson et al. (2014) found whale sharks during the years 2006-2010 were
most commonly encountered within 3 km of the reef front at the northern Ningaloo WSEI
area and within 6 km of the reef front at the southern WSEI area, it would be expected that
the NRETA would have a strong likelihood to record a tagged whale shark if moving within
the Ningaloo WSEI area overall. In fact, results from the current acoustic tracking study
indicates that whale sharks are most commonly recorded within 1 – 3 km of the reef front,
largely vindicating the Ningaloo WSEI practice of employing a spotter plane area search
within 5 km of the reef. However, a more even distribution of detections further offshore
from the reef crest at the Southern Line indicates that whale shark movements are not
restricted inshore when traversing the western edge of the mainland (i.e. near Point Edgar and
Point Cloates (see Figures 4.1 and 4.3)). In addition, whale sharks, although present, are
likely to go unreported if relying solely on WSEI monitoring, which does not search this far
offshore and in the non-tourism area north of Point Cloates and south of Yardie Creek.
Retention of external acoustic tags is difficult to assess as sharks are not available to
the observer (ecotourist) throughout the year at Ningaloo and therefore presence/absence of
an acoustic tag cannot be confirmed via photography. Long-term retention/tag loss may
present an issue and may not allow for a full 12 month data collection period, i.e. if these
sharks remain in the WSEI area at Ningaloo Reef throughout the year or if/when individuals
‘return’ to Ningaloo in ‘season’ subsequent to tagging. Despite this, one whale shark (A-815)
retained its acoustic tag for 11 months and was recorded within the Ningaloo Marine Park in
all but two of those months, providing almost year-round residency. It is possible however
that whale sharks do depart Ningaloo for an extended period, given both individuals
77
monitored over the greatest timescale, i.e. A-815 and A-411 (11 and 6 months, respectively)
were not recorded within NRETA for an extended period.
In another acoustic study on whale sharks by Cagua et al. (2015), tag retention rate
was less than 100%. Of 30 whale sharks tagged in Tanzania, 11 were confirmed (via visual
assessment) to have lost their acoustic tags over the course of the study (Oct 2012 to Nov
2014), with one shark having lost its tag after only 3 months (December 2012). However, the
total number of tagged individuals still retaining their acoustic tags some 12 months post-
deployment was 26.
Of the 21 individual whale sharks tagged within the NMP WSEI area, only seven (7)
individuals were recorded within NRETA on the day of tagging. In addition, despite photo-
identification confirmation of presence within the WSEI area (see Appendix 3), for two of the
three whale sharks where the acoustic tags had been removed prematurely in 2013, no
transmissions were received within the NRETA on the day of removal. It is therefore clear
that acoustic monitoring via NRETA does not capture all sharks all of the time. This may
indicate that the distribution of acoustic receivers within NRETA is spaced too far apart to
‘capture’ the presence of tagged whale sharks at Ningaloo on all occasions. As such, any
future assessment of whale shark residency at Ningaloo using acoustic data may benefit from
the deployment of additional receivers within NRETA, and those that are placed nearer to the
outer reef.
For the period 2011-2013, a relatively low residency index (RI) of 0.075 (as gauged
by photo-identification) compared with a RI of 0.18 (via acoustic data) for the same
individual acoustically tagged whale shark differed significantly, suggesting that despite
some individuals being recorded at Ningaloo for most months throughout the year (see Figure
4.4), either the whale sharks are not regularly available to photo-identification monitoring via
the WSEI or alternatively, the whale sharks are more likely to be simply moving through
78
Ningaloo Marine Park, rather than remaining for the calculated period of residency, with
some returning back at a later date. Alternatively, because the WSEI is seasonal, the majority
of tourism activities are restricted to a five-month period each year, in contrast to the
potential for year-round monitoring via NRETA should the whale sharks remain at Ningaloo
and the acoustic tags remain attached to the individual sharks. In addition, WSEI activities
are confined to restricted areas within Ningaloo Marine Park relative to coverage through
NRETA, therefore whale shark presence may be underrepresented.
While both photo-identification and acoustic monitoring have provided localised data
to better understand whale shark residency patterns within Ningaloo Marine Park,
presence/absence of whale sharks outside Ningaloo Marine Park could not be assessed as
acoustic detections are restricted by the number and location of the receivers within the Park,
while photo-identification is solely restricted to ecotourism operations that only occur within
the Park. To improve monitoring of this species within and beyond Ningaloo Marine Park,
the introduction of a satellite tracking program, an expanded acoustic tagging regime (to
include an increased number of sharks tagged; deployment of additional receivers within
NRETA; and improvements in technology to increase tag retention rate) and continuation of
the citizen science photo-identification reporting program should be considered in future
years.
79
Figure 4.1: Whale shark ecotourism operating region (South Ningaloo and North Ningaloo) of Ningaloo Marine Park, Western Australia.
80
Figure 4.2: a) VEMCO V16 acoustic tag with CETA tag anchor prior to deployment; b) V16
acoustic tag deployed on a whale shark at Ningaloo Marine Park.
81
Figure 4.3: Acoustic receivers deployed at Ningaloo Reef. (a) Encapsulates the Northern
Line, Mangrove Bay Array and Turquoise Line, (b) encompasses the Central Line between
Point Edgar and Point Cloates, and (c) the Southern Line near Coral Bay.
82
Month
Pro
porti
on (%
) of t
agge
d sh
arks
det
ecte
d
0
20
40
60
80
1002011 2012 2013
J F M A M J J A S O N D J F M A M J J A S O N D J F M A M J J A
WESI season
WESI season
WESI season
2011 2012 2013
Figure 4.4: Proportion of acoustically tagged whale sharks detected at Ningaloo Marine Park in 2011, 2012 and 2013.
83
Figure 4.5: Map of detections at acoustic receivers at Ningaloo Marine Park, Western
Australia.
84
Table 4.1: Distance (km) between acoustic receiver lines within the Ningaloo Reef
Ecosystem Tracking Array (NRETA) used to determine seasonal residency and distances
moved of acoustically tagged whale sharks.
General Description of acoustic installation
GPS at reef edge # receivers outside reef edge
Distance between receiver ‘lines’
Northern Line (NL) -21.8904, 113.9231 7 NL-MA = 10.1km
Mangrove Array (MA) -21.9790, 113.9015 24 MA-TL = 12.3km
Turquoise Line (TL) -22.0873, 113.8774 4 TL-CL = 62.5km
Central Line (CL) -22.5974, 113.6206 9 CL-23rd = 14.7km
23rd Parallel West (23rd) -22.9988, 113.7605 2 23rd-SL = 14km
South Line (SL) -23.1236, 113.7122 19
* Receivers are separated by approximately 0.8km within each line.
85
Table 4.2: Whale shark monitored via acoustic telemetry at Ningaloo Marine Park (2011-
2013).
Shark# Tag date Final detection
date Days
monitored Days
detected Residency
Index
A-029 20.09.11 12.10.11 23 2 0.09 A-087 05.05.13 24.05.13 20 7 0.35 A-101 09.09.11 02.11.11 55 9 0.16 A-108 11.09.11 02.12.11 83 14 0.17 A-236 10.04.13 23.04.13 14 5 0.36 A-273 22.05.12 29.05.12 8 2 0.25 A-411 13.09.11 09.02.12 150 16 0.11 A-546a 21.05.12 18.06.12 29 3 0.10 A-546b 09.04.13 19.05.13 41 13 0.32 A-579a 11.09.11 12.12.11 93 28 0.30 A-579b 15.06.13 06.07.13 22 5 0.23 A-584 15.09.11 09.10.11 25 3 0.12 A-749 19.06.13 24.08.13 67 9 0.13 A-783 22.05.12 01.10.12 133 12 0.09
*A-795 06.06.13 07.06.13 2 1 0.5* A-815 22.05.12 25.04.13 339 47 0.14 A-844 09.09.12 26.10.12 48 6 0.13 A-847 21.05.12 29.05.12 9 1 0.11 A-848 22.05.12 20.06.12 30 5 0.17 A-884 09.04.13 20.07.13 103 14 0.14
Mean
64.7 9.6 0.18 Minimum
2.00 1.00 0.09
Maximum
339.00 47.00 0.36 SE
±17.3 ±2.4 ±0.02
* Shark A-795 was excluded in calculation of residency index to reduce bias (only monitored for 2 days).
86
Table 4.3: Period of photo-identification for individual whale sharks involved in acoustic tagging study (2011-2013).
Shark # First detection (2011)
Final detection (2011)
Days mnt’d
Days det’d
First detection (2012)
Final detection (2012)
Days mnt’d
Days det’d
First detection (2013)
Final detection (2013)
Days mnt’d
Days det’d
RI
A-029 05.04.11 20.09.11 159 3 0.019 *A-087 05.05.13 05.05.13 1 1 A-101 22.05.11 09.09.11 111 4 0.036 A-108 01.06.11 11.09.11 103 2 0.019 A-236 09.04.13 23.04.13 15 4 0.267 *A-273 21.05.12 22.05.12 2 2 *A-411 13.09.11 13.09.11 1 1 A-546a 21.05.12 05.07.12 46 4 0.087 A-546b 09.04.13 04.06.13 57 2 0.035 *A-579a 09.09.11 09.09.11 1 1 A-579b 18.05.13 29.06.13 43 3 0.070 A-584 30.04.11 15.09.11 139 2 0.014 A-749 19.06.13 09.08.13 52 3 0.058 A-783 26.04.12 28.05.12 33 4 0.121 A-795 14.05.13 08.06.13 26 5 0.192 A-815 22.05.12 01.10.12 133 3 0.022 A-844 31.05.12 09.09.12 102 2 0.020 *A-847 21.05.12 21.05.12 1 1 *A-848 22.05.12 22.05.12 1 1 A-884 19.03.13 09.04.13 22 2 0.091 Mean 85.67 2.16 45.43 2.43 30.86 2.86 0.075 Min. 1 1 1 1 1 1 Max. 159 4 133 4 57 5 SE ±27.99 ±0.48 ±20.03 ±0.48 ±7.75 ±0.51 ±0.02 * For the Residency Index calculation, A-273 was excluded because monitoring was over only 2 days; A-087, A-411, A-579a, A-847 and A-848 were excluded as monitoring was for only a single day.
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Chapter 5
Norman BM, Reynolds S, Morgan DL (2016) Does the whale shark
aggregate along the Western Australian coastline beyond Ningaloo
Reef? Pacific Conservation Biology 22(1), 72-80.
Whale sharks (Rhincodon typus) seasonally aggregate at Western Australia’s Ningaloo Reef in
the austral autumn and winter, but their occurrence beyond this region during spring and summer
remains elusive. The aggregation at Ningaloo Reef coincides with a pulse of productivity
following mass coral spawning in early autumn, with the population during this period
dominated by juveniles that amass for feeding purposes. To investigate their movement patterns
beyond Ningaloo Reef, whale sharks were fitted with SPOT (n = 13) or SPLASH (n = 1) tags
between April and September (2010–14). Tagged whale sharks ranged in total length from 3 to 9
m. Each whale shark was also photographed for its subsequent identification using Wildbook for
Whale Sharks, and their years of residency at Ningaloo Reef determined. Temporal and spatial
observations of whale shark sightings were also determined through the conducting of interviews
with people throughout 14 coastal towns along the Western Australian coastline, as well as
through historical sightings and the Wildbook database. Satellite tracking revealed that all sharks
remained relatively close to the Western Australian coast, travelling a mean minimum distance
of 1667 (±316, s.e.) km. Public reports, coupled with satellite tracking, demonstrated that whale
sharks inhabit most of the Western Australian coast (from 35°S to 12°S), and that seasonal
88
migrations beyond Ningaloo Reef may be to the north or south and may similarly be associated
with areas of increased productivity.
Brief summary: The use of satellite tracking, citizen science and photo-identification
demonstrates that whale sharks that aggregate at Ningaloo Reef, Western Australia, migrate
north and/or south along the expansive Western Australian coast, before returning to Ningaloo in
the austral autumn.
Introduction
The whale shark (Rhincodon typus), which is listed as Vulnerable on the International Union for
the Conservation of Nature’s Red List (Norman 2005), is known to aggregate seasonally around
areas of increased productivity (Colman 1997; Heyman et al. 2001; Nelson and Eckert 2007;
Hobbs et al. 2009; Motta et al. 2010). While there is limited mixing of whale shark genetic
material between ocean basins (Castro et al. 2007; Schmidt et al. 2010; Sequeira et al. 2013), the
complete migratory routes taken by whale sharks have yet to be fully documented and remain
relatively poorly understood (Rowat and Brooks 2012).
In Western Australia, Ningaloo Marine Park, which encompasses the Ningaloo Reef,
represents an important aggregation site for whale sharks in the Southern Hemisphere, and this
occurrence has been well documented since the 1980s (Taylor 1994; Wilson et al. 2001). This
appears to be directly influenced by the mass coral spawning event that occurs on Ningaloo Reef
around early to mid-autumn of each year (Colman 1997). Satellite tracking has previously
revealed that several individual whale sharks have been recorded moving north and north-east
away from Ningaloo Reef following their seasonal aggregation (Wilson et al. 2006; Sleeman et
89
al. 2010), although the duration of each of these tracks was always less than 12 months. No
satellite telemetry study has, to date, been able to confirm evidence of a return to the feeding
aggregation at Ningaloo Reef once individuals have departed this region. Relatively few whale
shark sightings have been recorded from along the Western Australian coast outside of the
Ningaloo Reef, and there have been no targeted interview-based surveys undertaken in these
regions.
Implementation of a robust data-collection program via citizen science and culminating
in the collation of these data from disparate sources within the Wildbook for Whale Sharks
(founded as the ECOCEAN Whale Shark Photo-identification Library) (www.whaleshark.org)
has provided an ideal monitoring program for the species. This database is a globally and
regionally scoped online research platform for standardised capture–mark–recapture recordings
for the species. The unique system utilises the natural skin patterning on whale sharks (Norman
1999; Arzoumanian et al. 2005), and through the support of volunteers via this citizen science
program it has been possible to document sightings of individual whale sharks at Ningaloo Reef.
As of 31 December 2014, >1000 individual whale sharks have been identified at this aggregation
site (www.whaleshark.org). However, photo-identification has suggested that only one of these
sharks has been recorded in any other global jurisdiction. Here, satellite tracking, together with
publicly sourced reports and photo-identification is used to test the hypothesis that the whale
sharks at Ningaloo Reef may either remain in the area, or stay close to the Western Australian
coast, and return to Ningaloo Reef in the Austral autumn to feed.
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Materials and methods
Satellite tag deployment
Whale sharks were located using a spotter plane at the northern end of the Ningaloo Marine Park
and tags were deployed while a researcher was swimming alongside each animal. Each tagged
whale shark was photographed to enable identification using the Wildbook database and the sex
and estimated size of each individual were recorded, where possible, at the time of tagging.
Photographs were taken according to well established protocol (Arzoumanian et al. 2005), i.e.
lateral photograph of the unique spot pattern behind gill slits. In total, 14 whale sharks were
satellite tagged between 2010 and 2014. Tagging generally occurred towards the end of the
seasonal aggregation of whale sharks at Ningaloo Reef, i.e. between July and September, in an
effort to gather data on these fish following their movement away from the aggregation zone.
Tagging was undertaken according to Murdoch University Animal Ethics Permit No. W2402/11
and permit nos SF009184 and CE003919 from the Western Australian Department of
Environment and Conservation and the Western Australian Department of Parks and Wildlife,
respectively.
In 2010, Wildlife Computers satellite tags (1 × SPOT; 1 × SPLASH) were embedded in a
positively buoyant syntactic foam towed body (Fig. 1a) by Nikolai Liebsch (Customized Animal
Tracking Solutions (CATS)) and attached by a 2-m wire trace to a stainless steel anchor (dart)
that was deployed on the flank of each whale shark just below the first dorsal fin using a Woodie
1000 speargun. In 2012, 2013 and 2014, Wildlife Computers satellite (SPOT) tags were attached
to a negatively buoyant fin-mount clamp manufactured by CATS (Fig. 1b) that was secured to
the first dorsal fin of each shark (Fig. 1c, d) as it was evident that the first dorsal fin is clear of
the water surface while the shark actively feeds at dusk (see Fig. 1e). Each arm of the clamp had
two 1-cm spikes, which, despite not penetrating the skin, enabled the unit to remain attached to 91
the fin. A galvanic time release was included in the design in 2012 (see Gleiss et al. 2009), while
in 2013 and 2014 a corrodible section (dissimilar metals: clamp arm and spikes) was employed.
In each instance, the release period was estimated (from previous laboratory trials) to occur at a
maximum of six months from the deployment date, to ensure that the clamp would drop off from
the fin resulting in minimal long-term impact on the animal. An advantage of the negatively
buoyant fin-mount tag is that if the tag is dislodged from the shark, ARGOS will receive no
transmissions (cf. continuous transmissions from the towed body tag) while floating freely at the
surface despite being dislodged from the shark (see Gifford et al. 2007; Hearn et al. 2013). In
addition, the fin tag design is likely to induce less drag, demand less attention from predatory
fishes that may attack this unit (which in the towed body design, can resemble a lure), and
accumulate less weed and organic fouling that may impact tag buoyancy in the towed body.
To establish the physical impact of the tagging on individuals at Ningaloo Reef, when a
previously tagged (and photo-identified) shark was resighted, a photograph of the first dorsal fin
was collected where possible.
Satellite tags reported locational positions via the ARGOS satellite network to CLS
AUST-NZ-South Pacific and these were subsequently downloaded daily for this study. ARGOS
assigns a Location Class (LC) to each, with LC 3, 2, 1 and 0 when at least four uplinks are
received on an overpass; LC A is established from three uplinks; while LC B occurs when only
two uplinks are available (Hays et al. 2001). Position estimates had an associated error: LC of 3
(<250 m), 2 (250–500 m), 1 (500–1500 m), 0 (>1500 m), A and B (not specified) (www.argos-
system.org). Location Class A and B were still included in the analysis, providing these
positions were not of a distance so far removed compared with earlier and later more-accurate
positions that they were considered erroneous.
92
ARGOS data were converted to the format preferred by ZoaTrack (https://zoatrack.org/)
and subsequently uploaded to this public website, enabling the tracks of the tagged whale sharks
to be easily observed. ZoaTrack automatically calculates the straight-line distance between each
transmission location (irrespective of whether a land mass is present between two consecutive
locations) and combines these to establish minimum distance travelled. Erroneous positions are
removed to ensure these do not incorrectly influence distance calculations.
Historical and community sighting records
To enable the documentation of whale shark sightings at times and locations outside the regular
season (April–July) at Ningaloo Marine Park, sighting surveys were conducted from 405
attendees at public meetings during visits to 14 towns along ~3000 km of the Western Australian
coastline between Albany and Broome (see Fig. 2). Additional reports were submitted from the
general public for this study via the internet, by telephone, and during personal interviews
conducted with stakeholders involved with oil and gas facilities along and offshore throughout
Western Australia’s north-west coast.
Whale shark photo-identification
The Wildbook for Whale Sharks database provides the basis for a comprehensive data collection
regime for each annual whale shark season at Ningaloo Reef. The authors and the Western
Australian Department of Parks and Wildlife collect sighting data and identification images from
the Ningaloo Whale Shark Ecotourism Industry Licensees and staff. Volunteers coordinated by
the authors then process the appropriate images according to Arzoumanian et al. (2005). Using
computer-assisted scanning technology, it is possible to determine whether the individual whale
93
shark in question is a ‘new’ shark or a ‘resight’ of a previously reported whale shark within the
database. Sighting data from Ningaloo Marine Park has provided the longest continuous dataset
of global whale shark encounters, between 1995 and 2014, and has enabled the determination of
the number and frequency of returning individual whale sharks to this location.
Results
Satellite tagging
Of the 14 whale sharks fitted with satellite tags between July 2010 and June 2014, 13
successfully transmitted positional data, with the number of days from tag deployment to last
transmission ranging from 9 to 261 (mean = 78.5 (±20.5, s.e.)) (Table 1). Nine males and four
females were tagged, with one fish unable to be sexed at the time of tag deployment. The males
ranged in total length (TL) from 3 to 7 m (mean TL = 5.4 (±0.47, s.e.)), while the females ranged
in TL from 5.5 to 9 m (mean TL = 6.75 (±0.64, s.e.)). The mean minimum distance that these
individuals travelled was 1667 (±316, s.e.) km, and ranged from 110 to 3214 km. All tagged
individuals remained in the approximate vicinity of the Western Australian coastline from the
Timor Sea in the north off the Kimberley coast, to waters some 1000 km north-west of Ningaloo
Reef, to the south-western corner of the Western Australian capital city of Perth (Fig. 3) –
essentially between 12° and 35°S. Two of the male whale sharks (A-683 and A-660) moved in
an east-north-east direction, as far north as the Timor Sea, and off the coastal Kimberley town of
Broome.
Eight days after tag deployment in 2013, shark A-720 was recorded at Shark Bay, before
returning to Ningaloo Reef two weeks later. Over the subsequent month, A-720 was recorded
moving in a southerly direction and remained at Shark Bay from mid-October until final
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transmissions were received on 7 November. A-660 and A-843 were also tracked to Shark Bay
in 2010 and 2012, respectively.
Of the four female sharks tagged, the tag on one transmitted for only nine days. The
remaining three were tracked for between 74 and 92 days, travelling between 1653 and 3123 km,
and all moved west-north-west from Ningaloo Reef, at distances of 1–1000 km from the coast,
two in a depth of up to 5000 m. Shark A-546 (a male) was tracked for 261 days, remaining close
to Ningaloo Reef for most of the track. A further four sharks, three of which could be confirmed
as male and the fourth unknown (A-660, A-481, A-302, A-843) were tracked to near the oil and
gas facilities ~140 km north-west of Karratha.
Shark A-633 was tracked for 203 days over a distance of 3214 km, first moving 600 km
north-west of Ningaloo Reef before returning in November, then moving south until additional
transmissions were received ~200 km west-north-west of Perth in early February (Location
Class 1). Shark A-633 was last recorded (Location Class 1) in early March ~50 km west of its
previous position, north-west of Perth in a water depth exceeding 5000 m.
Citizen Science sighting reports
More than 8000 sightings were recorded of whale sharks along much of the Western Australian
coastline (see Fig. 2). Whale shark sightings from Western Australia were reported throughout
the calendar year, with some areas showing increased frequency in particular months (Fig. 4).
For example, while the Ningaloo Marine Park had an extremely high number of sightings
between April and July, the region immediately north (i.e. Barrow Island/Karratha) had peak
sightings in August, while the area immediately to the south of Ningaloo recorded peak sightings
between December and March. Further to the south (Lancelin to Albany), sightings were
95
generally restricted to summer months and early autumn. Occasional sightings in the Kimberley
were restricted to between July and December.
Photo-identification
Several additional whale sharks were positively identified outside of the Ningaloo Reef using the
photo-identification program. For example, one whale shark was filmed by staff of an oil and
gas facility (off Barrow Island) at a location 250 km from Ningaloo Reef during 2008, and
subsequently confirmed through photo-identification as A-504. This individual was also sighted
at Ningaloo Reef in 2006 and 2009 using photo-identification. A second shark (A-535) was
sighted ~500 km north-east of Ningaloo Reef (from the North West Shelf Nan Hai 6 facility) in
2005, and at Ningaloo Reef in all years between 2009 and 2012. Shark A-624 was photographed
on 7 October 2009, 120 km north of Karratha (~350 km from Ningaloo Reef) and resighted at
Ningaloo Reef the following year on 10 April 2010. The photo-identification program also
confirmed the identity of two whale sharks at Shark Bay, one of which was recorded at Ningaloo
in June 2001 (A-230), at Shark Bay in May 2004, and at Ningaloo in June 2005. Another (A-
290) was recorded at Ningaloo Reef in June 2002 and at Shark Bay in May 2004.
Shark A-633 was tracked for 203 days. During this time, the shark had moved from
Ningaloo to 500 km north-west of Barrow Island and then a further 1500 km south to a location
west of Rottnest Island. This individual was resighted back at Ningaloo in 2015 and represents
the longest confirmed movement away from, and returning to, Ningaloo to date. The minimum
distance covered exceeds 3000 km.
Using photo-identification, only one whale shark from beyond the Western Australian
coast (or at least closer to the coast of another jurisdiction) has been confirmed in Western
96
Australian waters: a Whale Shark (A-424) photographed off the coast of Borneo in 2007 and
submitted to the Wildbook database by a member of the general public in 2009 was positively
identified from Ningaloo Marine Park in 2011 and 2012. The minimum straight-line distance
between both sites exceeds 2500 km (although likely greater as a land mass between must be
negotiated) and represents one of the greatest recorded one-way movements across hemispheres
for this species.
Where images of the first dorsal fin of a previously tagged shark were available, the
degree of injury attributable to tagging was negligible on each occasion (see Fig. 5).
Discussion
Photo-identification of whale sharks at Ningaloo Reef has provided a great deal of information
relating to mark–recapture at this globally important feeding aggregation (Norman 1999;
Holmberg et al. 2008, 2009). However, much of this information is limited spatially and
temporally – specifically to the annual aggregations at Ningaloo Reef and the Whale Shark
‘season’, which extends throughout the austral autumn and winter (Colman 1997). Beyond this
spatial and temporal scale, little information is available on the occurrence and movement
patterns of whale sharks throughout the expansive Western Australian coastline, which is one of
the most sparsely populated coasts on the planet. This study explored the spatial and temporal
patterns of whale shark occurrences throughout ~3000 km of the Western Australian coast, using
satellite tracking, photo-identification and the collation of sighting data from the general public
to demonstrate that whale sharks are found throughout Western Australia’s coastal and offshore
Indian Ocean waters from the Timor Sea (Kimberley) in the north, south through the Pilbara
coast, to Ningaloo, Shark Bay, the Mid-west and south to the Southern Ocean waters off Albany.
97
These data complement previous studies from Ningaloo Reef, including results from 19
individuals satellite-tagged there between 2002 and 2005 showing that whale shark distribution
from Ningaloo is predominantly ia a north-west, north and north-east direction (Wilson et al.
2006; Sleeman et al. 2010); however, there is no photo-ID record of these individuals in the
database, so it is not possible to determine in which years these animals may have returned to
Ningaloo. One individual (identified as A-100 – see
http://www.whaleshark.org/individuals.jsp?langCode = en&number = A-100) satellite tagged in
2002 moved more than 2500 km towards Christmas Island, Indian Ocean, before shedding its tag
after ~45 days (J. Stevens, unpubl. data). A second whale shark also tagged in 2002 (and
subsequently identified as A-112) was tracked moving north-north-east away from Ningaloo
Marine Park towards Indonesia (a distance travelled of >1500 km) before prematurely shedding
its tag some 35 days later (J. Stevens, unpubl. data). Photo-identification data revealed that A-
112 was subsequently recorded at Ningaloo Reef in 2005, 2007, 2008, 2009, 2010 and 2013 (see
http://www.whaleshark.org/individuals.jsp?number = A-112&langCode = en&Go = Search).
Although useful for providing important locational information when individuals are
away from human sight, satellite-tagging of whale sharks in particular provides challenges,
especially compared with satellite tagging of air-breathing animals, i.e. transmission is possible
only when the tag is dry and the antenna free of the ocean surface (Acuña-Marrero et al. 2014).
The limitation of satellite tags, other than becoming dislodged due to poor attachment methods,
may be restricted to periods when whale sharks feed at the surface, which may be for only ~8
min per day (see Gleiss et al. 2013). Nonetheless, the results from the current study may have
revealed new foraging grounds for this large filter-feeding shark. For example, the three female
whale sharks (A-349, A-666, A-481) with deployments longer than nine days were moving
through the only known spawning ground for southern bluefin tuna (Thunnus maccoyii) (Caton
1991) at the time when spawning is known to occur, i.e. August–April, with a peak in October
98
(Honda et al. 2010). Shark A-633 was satellite-tracked further south than any previously
identified whale shark, to an area west of Perth (near the Rottnest Trench), where large
aggregations of samson fish (Seriola hippos) are known to spawn between November and
February (Rowland 2009). This individual was recorded in two locations less than 50 km apart
in early February and again in early March. It thus seems plausible that whale sharks could also
be targeting areas of high productivity for opportunistic feeding on energy-rich resources, as
they are known to do elsewhere, including near tuna spawning grounds in the Coral Sea,
Australia (Colman 1997), at a snapper (Lutjanus sp.) spawning site off Belize (Heyman et al.
2001; Graham et al. 2006), and near rich plankton blooms off the Yucatan Peninsula, Mexico
(Motta et al. 2010).
Although historically whale sharks are known from Ningaloo Reef over a particular
‘season’ (Colman 1997), the data presented confirms that whale sharks are commonly present at
this locality outside this restricted period. Whale shark sightings may be low in the non-tourist
season because of the remoteness of the location and the lack of a tourist industry outside April–
July at Ningaloo Reef. Waters to the south of Ningaloo Reef, between Shark Bay and Geraldton,
were found to be an area with the highest whale shark sightings outside of Ningaloo Marine
Park, predominantly between October and March. Here, it is likely that the westerly winds and
ocean currents concentrate late larval-stage (puerulis) western rock lobsters (Panulirus cygnus)
settling on inshore reefs along the west coast mainly between August and January each year
(Caputi et al. 2001), thus providing increased feeding opportunities for plankton-feeding whale
sharks at this time.
Further north-east from Ningaloo Reef, citizen science sighting reports from workers at
oil and gas facilities and/or participants in humpback whale (Megaptera novaeangliae) survey
flights provide evidence for whale shark presence around oil and gas facilities. Photo-
99
identification submissions to the Wildbook database from subsea remote-operated vehicle
footage has recorded two whale sharks around oil and gas facilities at depths exceeding 100 m,
where they were feeding on small fish aggregating around these structures. And of the 13 sharks
for which we have satellite data, four (A-660, A-843, A-481, A-302) were recorded surfacing
near oil and gas facilities close to the Goodwyn and Rankin fields (~140 km north-west of
Karratha). This lends support to the possibility that these facilities provide a type of fish
aggregation device for whale sharks or, more specifically, their prey. Robinson et al. (2013)
report on whale sharks aggregating at oil and gas platforms ~90 km offshore in Qatari waters,
suggesting that these are effectively human-made offshore reefs that support an increased
biodiversity compared with waters away from the platforms. It seems possible that oil and gas
facilities therefore could have broader consequences: the natural migration route of whale sharks
and the timing of such may be affected by the presence of these non-natural structures in the
region.
Of the 14 whale sharks satellite-tagged at Ningaloo Reef, 13 were recorded at this
locality via photo-identification in years other than the year of tagging, with four photographed
in each year between 2010 and 2014, five in each year between 2010 and 2013, and five of the
tagged sharks sighted again in 2015. It is possible that the shark sighted in only a single year (A-
883) was either a new recruit to the population or it may not have yet been recorded during an
ecotourism interaction, providing the opportunity to confirm its presence by photo-identification.
The high seasonal site fidelity for most whale sharks suggests strongly that they return to
Ningaloo Reef over several years, which has also been recognised in previous photo-monitoring
studies (Arzoumanian et al. 2005; Holmberg et al. 2008, 2009). Although Ningaloo Reef is a hot
spot for this species, with individuals often sighted there between April and July (the general
whale shark tourist season), coinciding with a period of high productivity (Sleeman et al. 2010),
results from the current study show individuals to distribute broadly from this location off the
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Western Australian coastline, with most migrating to the south between Shark Bay and
Geraldton. This suggests that the animals search independently for, and exploit, other feeding
opportunities while regularly returning to Ningaloo Reef where the feeding resource is consistent
around the time of the annual mass coral spawning and the months that follow.
While tourism-derived photographic data have been able to provide evidence of inter-
aggregational migration between closely located countries (e.g. Mexico to USA to Belize and
Honduras; South Africa to Mozambique, Mozambique to Tanzania), these ‘movements’ were
over short distances (i.e. ≤1000 km) (Norman, unpubl. data). In contrast, the movement of A-424
over ~2500 km is rare. It does, however, indicate the possibility of recruitment of whale sharks
from Indonesian waters to Australia, which may account for limited genetic mixing between
jurisdictions (see Castro et al. 2007; Schmidt et al. 2010; Sequeira et al. 2013).
These results verify that whale sharks are distributed further along the Western
Australian coast than just at Ningaloo Reef, and that these individuals appear to aggregate
seasonally at Ningaloo Reef. The perceived ‘seasonality’ in this species is not entirely accurate,
however, as this study provided evidence of year-round sightings of whale sharks at Ningaloo
Reef. Other regions beyond Ningaloo Reef may provide important feeding opportunities for
whale sharks, including between Shark Bay and Geraldton (western rock lobster peurulis), south
of Java (southern bluefin tuna spawn), and off Rottnest Island (samson fish spawn). In addition,
oil and gas facilities could potentially affect the movement patterns of whale sharks by providing
feeding opportunities exploited by this species. Despite the track of A-546 (261 days)
representing the longest duration for a satellite-tracked whale shark from Ningaloo, it is still
short of a full year and the complete annual migration of the species remains elusive. However,
the results presented in this study demonstrate the benefit of combining citizen science with
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minimal-impact electronic tracking studies to determine whale shark movements over an
extended timeframe.
Acknowledgements
Our thanks go out to the providers of data for this study, including the people of Western
Australia in the towns along the coast. We especially thank Caroline Bayer, Sandra Leigh,
Western Australian Department of Parks and Wildlife (DPaW), ECOCEAN Australia,
Department of Fisheries Western Australia, Whale Shark industry stakeholders at Ningaloo
Marine Park, Western Australia, offshore oil and gas employees, Pam and Paul Dickenson, and
Curt and Micheline Jenner. This research has made use of data and software tools provided by
Wildbook for Whale Sharks, an online mark–recapture database operated by the non-profit
scientific organisation Wild Me with support from public donations and the Qatar Whale Shark
Research Project. Additional data entry and preparation of figures was undertaken by Darcy
Bradley, Briana Canny, Andrew Needham and Dr Stephen Beatty, for which we are indebted.
Special thanks to Kim Hands and Kylie Maguire for their work in gathering community data
between Albany and Broome in 2013, supported through The Garry White Foundation,
Mitsubishi Australia and the Inspiring Australia Initiative (a joint project between Scitech, the
State and Federal Government). We appreciate the assistance of Guan Oon and Hollie Lourie
(CLS AUST-NZ-South Pacific) with satellite data analysis and acknowledge the satellite data
visualisation portal provided by ZoaTrack. Satellite tags were either funded or donated by Mr
Steve Wall, Novotel Ningaloo Resort, Hearts for Sharks, Driftwood Jewellers, Murdoch
University, DPaW, WA Department of Tourism, Australian Department of the Environment,
CATS and ECOCEAN. We thank Nikolai Liebsch (CATS) for his refinement in the design and
building of the tag attachments used in this study. We also thank Adrian Gleiss and Nikolai
102
Liebsch for insightful comments on an earlier version of the manuscript. The work was
conducted according to Murdoch University Animal Ethics Permit No. W2402/11, and DPaW
Permits Nos SF009184 and CE003919. Author Contributions: conceived and designed the
experiments, BN, DLM; analysed the data, BN, SR, DLM; contributed
reagents/materials/analysis tools, BN, SR; wrote the paper: BN, DLM.
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Figure captions
Fig. 1. SPOT tag embedded in: (a) towed body and using the (b) fin clamp design; (c) fin clamp tag being deployed; (d) fin clamp tag attached to first dorsal fin; (e) fin clear of water surface while feeding at dusk at Ningaloo Marine Park. Photographs: Brad Norman.
Fig. 2. Sightings of whale sharks from submitted reports and personal interviews with ocean users along the Western Australian coastline.
Fig. 3. (a) Combined tracks of 13 whale sharks that were tagged at Ningaloo Reef between 2010 and 2013. Satellite track of female whale sharks (b) A-481, (c) A-349, and (d) A-666 satellite tracked after leaving Ningaloo Marine Park.
Fig. 4. Regional whale shark sightings (north to south) recorded during this study in particular regions in Western Australia between 2000 and 2014. See Fig. 2 for locality details.
Fig. 5. Examples of whale sharks resighted in years subsequent to tagging showing minimal impact caused from tag clamp.
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Fig. 1.
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Fig. 2.
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Fig. 3.
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Kimberley
Month
Jan
Feb Mar AprMay Ju
n Jul
Aug Sep Oct Nov Dec
0
25
50
75
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Pilbara
Month
Jan
Feb Mar AprMay Ju
n Jul
Aug Sep Oct Nov Dec
0
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50
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Ningaloo
Month
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Feb Mar AprMay Ju
n Jul
Aug Sep Oct Nov Dec
Num
ber o
f Wha
le S
hark
s
0
500
1000
1500
2000
2500
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MId-west
Month
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Feb Mar AprMay Ju
n Jul
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25
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South-west
Month
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Feb Mar AprMay Ju
n Jul
Aug Sep Oct Nov Dec
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Fig. 4.
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Fig. 5.
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Table 1. Details of satellite tag deployments on whale sharks at Ningaloo Marine Park between 2010 and 2014
Shark Sex Length (TL) (m)
Date tag deployed
Date of last transmission
Minimum No. days tag remained attached
Minimum distance travelled (km)
Years recorded by photo-identification at NMP
A-683 M 6 16.07.10 29.08.10 45 1906 2010, 2011, 2012
A-660 M 6 16.07.10 20.09.10 67 2571 2010, 2011, 2012, 2013
A-843 N/A 8 09.09.12 21.10.12 43 1183 2008, 2012, 2013
A-844 M 3.5 09.09.12 - - - 2010, 2012, 2013
A-546 M 7 09.04.13 25.12.13 261 3165 2009, 2011, 2012, 2013, 2015
A-633 M 5 24.08.13 14.03.14 203 3214 2008, 2010, 2011, 2013, 2015
A-720 M 5.5 21.08.13 07.11.13 79 1328 2010, 2012, 2013
A-481 F 6.5 07.07.13 06.10.13 92 1653 2007, 2008, 2009, 2010, 2011, 2012, 2013, 2014, 2015
A-666 F 5.5 20.07.13 05.10.13 78 2227 2010, 2011, 2012, 2013, 2015
A-349 F 9 03.08.13 15.10.13 74 3123 2004, 2005, 2008, 2009, 2010, 2011, 2013
A-534 M 6.5 16.06.13 04.07.13 19 110 2009, 2010, 2011, 2013, 2014, 2015
A-883 M 3 10.04.13 22.04.13 13 228 2013
A-088 F 7.5 21.06.14 29.06.14 9 358 2003, 2008, 2010, 2011, 2012, 2013, 2014
A-302 M 5 21.06.14 27.07.14 37 601 2006, 2007, 2009, 2010, 2011, 2012, 2013, 2014, 2015
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Chapter 6 General Conclusions
The use of a combination of invasive and non-invasive techniques during this study has
allowed the determination of a number of key ecological conclusions to be drawn of local
(Western Australia) and global aggregations of whale sharks, a species that is listed as
vulnerable by the IUCN. Gathering data on the life of this enigmatic species remains a
challenge, but the combination of scientific studies and the inclusion of data derived from
citizen scientists have allowed the collection of data that has provided regional differences in
population dynamics and migration patterns. Chiarucci et al. (2011) note that the measurement
of biogeographic and biological data over large geographic areas is simply not feasible by a
single team of researchers, and the whale shark proved to be the ideal candidate species to
support this notion.
The combination of citizen science and telemetry studies has, importantly, from a local
perspective, found that whale sharks in Western Australia seasonally congregate at Ningaloo
Reef in the Austral autumn and winter in large numbers, and then remain in this jurisdiction for
many years. From here, over 1100 individuals have been identified using observations from
scientists and the general public. Further, citizen science has allowed knowledge to be
generated on the extent of occurrence of the species in Western Australia, where they have
been reported from over 3000 km of coastline. Satellite tracking of a selection of individuals
has supported these data. However, although the aggregation at Ningaloo appears to be
concentrated to the Austral autumn and winter, telemetry studies have determined that a
number of individuals are found at Ningaloo for extensive periods outside of the whale shark
‘season’.
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Although satellite tracking has revealed that individuals within Western Australia may
migrate north towards Indonesia, only one individual has ever been identified in these different
jurisdictions using photo-identification. This study is the first to determine that the Western
Australian population of whale sharks should be considered as a local population, based on the
evidence presented by telemetry and citizen reports that suggest that outside of the aggregation
period, individuals not only roam north and south along the coast, but that many do remain at
Ningaloo Reef, while some individuals appear to be feeding in other areas of high productivity.
Similarly, on a global perspective, trans-oceanic migrations of whale sharks appears unlikely.
An extraordinary long-term site fidelity among whale sharks at multiple global hotspots (up to
21 years at Ningaloo Reef, Western Australia, for example) is occurring, with many identified
whale sharks within these feeding aggregations returning to the same location in subsequent
years. This represents some of the longest monitoring of any individual fish, and in time may
allow monitoring to last for many decades. For the top 20 global hotspots, the fact that
approximately one third of all whale sharks return to a familiar site in a subsequent year(s)
indicates strong site fidelity in this species, and at Mozambique, Maldives and Honduras, there
is clear evidence of year-round whale shark appearance.
In the case of R. typus, the predominance of small and immature individuals evident at
most constellations studied appears to coincide with important regular natural feeding
opportunities, although that prey items are somewhat varied between constellation sites close
to the relative safety of a coastal environment (Clarke and Nelson 1997, Norman 1999,
Heyman et al. 2001, Jarman and Wilson 2004, Hoffmayer et al. 2007, Nelson and Eckert 2007,
Meekan et al. 2009, de la Parra Venagas et al. 2013, Fox et al. 2013, Gleiss et al. 2013,
Robinson et al. 2013, Rohner et al. 2013). While individual R. typus are small and immature,
the prime directive for members of these constellations may be to expend minimal effort to
find food and increase in size and relative fitness (especially to avoid predation) prior to
114
expending greater energy reserves in the search for mates and reproduction. This may be
achieved by exploiting shallower coastal aggregations of prey. Exactly where the individuals
reside for the remainder of the year remains largely undefined, although it is possible that
whale sharks are present but simply unavailable for capture by photo-identification monitoring
techniques. In addition, it is possible that larger individuals may have an increased ability to
forage deeper into the epipelagic and mesopelagic zones (Wilson et al. 2006).
Despite the apparent level of site fidelity evident in this study, a limited number of
individuals have been confirmed moving between one or more nearby countries via: photo-
identification e.g. Seychelles/Mozambique (Gifford 1998, Rowat and Gore 2007);
Belize/Mexico/Honduras/USA (McKinney et al. 2013); and satellite tracking studies e.g.
Taiwan/Japan/Philippines (Hsu pers. comm.); Australia/Indonesia (Sleeman et al. 2010);
Mozambique/Madagascar (Brunnschweiler et al. 2009); Utila/Belize/Mexico (Gifford et al.
2007); Mexico/ Saint Peter and Saint Paul Archipelago, Atlantic Ocean, Mexico/Cuba (Hueter
et al. 2013); and Saudi Arabia/Egypt/Yemen/Oman (Berumen et al. 2014). However, most of
these movements are relatively small and although it is commonly accepted that R. typus is
highly migratory (IUCN SSG/CMS 2007), few reliable records exist for extensive movements
across ocean basins (Hueter et al. 2013), and this study supports that.
The current study has confirmed that at least some individuals within whale shark
constellations undertake longitudinal movements, albeit at the largely sub-adult life stage, and
usually at coastal margins. Given favourable prey availability at each location (Sleeman et al.
2010, Rowat and Brooks 2012), these movements are potentially driven by feeding
opportunities.
Despite more than 6000 individual whale sharks identified at coastal hotspots
worldwide from data supplied from 4000+ individual researchers and citizen scientists and
collated within the ECOCEAN (Wildbook) Whale Shark Photo-identification Library, there
115
remains no matched sharks between different continents. It therefore seems unlikely that the
concept of a broad movement of coastal (young and immature) whale sharks occurs. Rather, it
is likely that prior to the onset of maturity, whale sharks take advantage of coastal feeding
opportunities, and then as they mature, they engage in more extensive migrations from each
population/s while still remaining within their native ocean basin as suggested within a recent
genetic study (Vignaud et al. 2014). Because of the paucity of large, mature individuals present
at these coastal constellations, opportunities to investigate such movements via photo-
identification or satellite tracking is extremely limited. Notwithstanding, the present study
using photo-identification does demonstrate linked connectivity between a number of coastal
constellation sites.
A key factor for sustainable management of whale shark/human interactions is a clear
understanding of the population dynamics of whale sharks, including intra- and inter-annual
variability in abundance and distribution (Colman 1997). Holmberg et al. (2008, 2009)
analysed population structure of whale sharks at Ningaloo, with indications that whale shark
numbers at North Ningaloo (where monitoring was greatest) showing positive growth,
indicating well-managed ecotourism is not having a negative effect on whale shark
appearance/numbers. More recently, the population trends of the species at Ningaloo appear to
be on the increase. Updated population assessments at other aggregation sites throughout the
world will in time provide an improved global understanding on the whale shark numbers. At
present, R. typus remains listed as ‘Vulnerable to Extinction’ in the most recent IUCN Red List
of Threatened Species assessment (Pierce and Norman 2016). However, of continued concern
for the future conservation of the species is hunting at a commercial scale (Li et al. 2012).
116
117
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Appendix 1
United Nations Environment Programme ECOCEAN Whale Shark Photo-identification (Field
Station) Manual [Holmberg J and Norman B (2009) ECOCEAN Whale Shark Photo-
identification - UNEP MANUAL. Technical Report (United Nations Environment Program -
Regional Seas) 69pp].
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Whale Shark Photo-Identification Library
© Kurt Amsler, ROLEX
FIELD STATION MANUAL
Acknowledgements 135
This ‘Manual’ has been produced with the support of the United Nations Environment Programme (UNEP) Regional Seas. Special thanks to Jason Holmberg for text and images and to Darcy Bradley for assistance with formatting and final edit.
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ECOCEAN FIELD STATION MANUAL
Table of Contents
OVERVIEW 128 1 MANAGING ENCOUNTERS AND SHARKS 129
1.1 BASIC ASSUMPTIONS 129
1.2 PROCESSING A NEW ENCOUNTER: APPROVE OR REJECT 131 1.2.1 REVIEWING A NEW ENCOUNTER 132 1.2.2 REJECTING AN ENCOUNTER 132 1.2.3 REACCEPTING A DATA-ONLY ENCOUNTER 132
2 EXTRACTING SPOT PATTERNS 133
2.1 BASIC ASSUMPTIONS 133
2.2 PRE-PROCESSING A PHOTO FOR PATTERN RECOGNITION 134 2.2.1 OBTAINING A SOURCE IMAGE FROM THE ECOCEAN LIBRARY 135 2.2.2 PRE-PROCESSING AN IMAGE IN PAINT.NET 136
2.3 SENDING A PATTERN TO THE ECOCEAN LIBRARY INTERCONNECT CLIENT 141
3 COMPARING SPOT PATTERNS WITH SHARKGRID 146
3.1 STARTING A SPOT PATTERN COMPARISON SCAN ON SHARKGRID 146
3.2 EXAMINING THE RESULTS OF A SCANTASK 147 3.2.1 PENDING SCANTASKS 147 3.2.2 COMPLETED SCANTASKS 148 3.3 ADMINISTERING SHARKGRID 148 3.3.1 THE SHARKGRID QUEUE: ADDING AND DELETING SCAN TASKS 148 3.3.2 VIEWING CONNECTED CLIENT NODES AND THEIR PERFORMANCE 148 3.3.3 CONFIGURING SHARKGRID BEHAVIOUR 149 4 INTERPRETING PATTERN MATCH RESULTS 151 4.1 MODIFIED GROTH RESULTS 151 4.1.1 METRIC RESULTS 151 4.1.2 VISUAL RESULTS 152 4.2 I3S RESULTS 153 4.2.1 METRIC RESULTS 153 4.2.2 VISUAL RESULTS 154
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5 MATCHING SHARKS 5.1 CREATING A NEW SHARK 5.2 ADDING AN ENCOUNTER TO AN EXISTING SHARK
155 155 157
6 SEARCHING THROUGH THE ECOCEAN LIBRARY 6.1 ENCOUNTER SEARCH 6.1.1 DATA EXPORT
157 158 158
FAQS 160 APPENDIX 1A HOW TO JOIN SHARKGRID 162
1A.1 DOWNLOADING THE SHARKGRID CLIENT 163
1A.2 INSTALLING AND STARTING THE SHARKGRID 163 1A.2.1 Windows 163 1A.2.2 Linux 164
APPENDIX 1B PHOTO KEYWORDS IN THE ECOCEAN LIBRARY 166 APPENDIX 1C SPOT! 177 1C.1 SPOT! REQUIREMENTS 178 1C.2 LOADING SPOT! 178 1C.3 BASIC SPOT! INSTRUCTIONS 178 1C.4 SPOT! FAQ 179 APPENDIX 1D. TAPIRLINK AND THE ECOCEAN LIBRARY 181 APPENDIX 1E. ECOCEAN LIBRARY LOCATION CODES 182 APPENDIX 1F. ECOCEAN LIBRARY ACCESS POLICY 183 APPENDIX 1G. ECOCEAN LIBRARY VISITOR AGREEMENT 185 APPENDIX 1H. USER AGREEMENT ‘ECOCEAN LIBRARY’ 189
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Overview
Welcome to the ECOCEAN Whale Shark Photo-identification Library User Community!
Overview of the ECOCEAN Library
The ECOCEAN Whale Shark Photo-identification Library is an Internet-based software application for cooperative whale shark research. The primary purpose of the Library is to increase our understanding of whale sharks on a global and local level and to promote related conservation efforts through high quality research and scholarship. On a purely functional level, the Library is used to collect, protect, store, and share whale shark mark-recapture data gathered from a variety of individuals and institutions worldwide.
Purpose of this Field Station Manual The purpose of this field manual is to document the standard operating procedures for the ECOCEAN Library that are required to ensure proper and consistent data collection, processing, and analysis. The intended audience for this document includes:
• current ECOCEAN Library users managing whale shark encounter data
• prospective Library users seeking to understand how the Library operates
• external reviewers seeking an in-depth look at Library operations
For more information about this manual, please contact: [email protected]
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1 Managing Encounters and Sharks
1.1 Basic assumptions The ECOCEAN Library is a mark-recapture framework for storing and analyzing whale shark data. As such, it divides data into two distinct types.
Encounters are individual sightings of whale sharks. An encounter report is submitted to the ECOCEAN Library via the Internet and represents a “mark” (first sighting) or “recapture” (subsequent re-sighting) of an individual whale shark. Each encounter contains photos and data that represent one whale shark at one point in time. An encounter can be added to a previously identified shark in the database, representing a resighting of that animal, or it can be allocated as a new individual shark and given a name, representing a new animal previously undocumented in the ECOCEAN Library. Encounters may also remain “Unassigned,” indicating that the encounter does not contain enough data to be identified as a new or previously seen shark at the current time, though it may be matched to other encounters in the future.
Sharks are uniquely identified animals and are made up of one or more encounters. To be assigned as a new shark, a new encounter must have a properly-oriented left-side spot pattern extracted from a photograph and added into the database. Encounters without a suitable left-side pattern can be added to an existing shark if some other characteristic (visual recognition of its pattern or of significant scarring, right-side pattern matching to another identified shark, etc.) can be used to link it to an existing shark in the Library. As we build up more and more encounters for each shark, we will be able to build up robust metrics for population analysis, allowing us to better understand whale shark biology and population trends on a local and a global scale.
Figure 1. One or more encounters make up an identified shark.
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1.2 Processing a new encounter: Approve or Reject The following steps describe how to approve or reject a newly submitted encounter for inclusion in the ECOCEAN Library. Sometimes encounters are added with incomplete or inaccurate information. It is best to leave these encounters as “Unapproved” until the submitter has been contacted and the information has been corrected.
1.2.1 Reviewing a new encounter
To review a new, unapproved encounter for inclusion in the library, follow these steps:
1. Look at the new encounter in the View Unapproved Encounters view (login required) and ensure that the thumbnail image was correctly rendered. If not, use the Reset thumbnail command on the encounter page to choose a photo to render a new thumbnail from.
2. Check to make sure that all of the photos submitted contain material appropriate for display on the ECOCEAN web site. This is an important check to make sure that offensive material or photos showing inappropriate interaction with whale sharks are not displayed in the library. If the photos contain appropriate content and have enough information (scars, visible left- or right-side patterns in photos, or extractable spot patterns) to reasonably expect that the shark can be re-identified now or in the future, proceed to the next step. If the encounter contains offensive material, no photos, or poor quality photos, proceed to “Rejecting an Encounter” below.
3. Click on each photo one by one to view it in full detail. Note the presence of any distinguishing characteristics, such as scars, distinct spot patterning, or physical tags, using the pulldown list of keywords above each photo.
4. Check and edit the data submitted for each encounter as needed. For example, verify that the size estimate is realistic (30 feet versus 30 meters) and that the submitted comments contain appropriate language and good spelling.
5. Add the appropriate Location Code to the encounter and check any reported GPS coordinates for accuracy using the displayed Google Map.
6. If the content of the unapproved encounter is acceptable, click Approve to make the encounter visible to the general public once all of the above steps have been completed.
7. Check the TapirLink status of the encounter (true/false). The ECOCEAN Library
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TapirLink policy is available here.
1.2.2 Rejecting an encounter
There are two reasons to reject an encounter:
1. The encounter does not contain enough information to uniquely identify the shark now or in the future, but it does contain some useful data such as date and location, sex, size, etc. that can later be used for trending and aggregate analysis.
2. The encounter contains offensive or inappropriate content. These encounters should be deleted permanently
To reject an encounter, click the Reject button in the left-hand blue bar of the encounter. This will give you two options:
1. Click Save as DATA-ONLY to preserve this encounter but not make it available
to the general public. Note: Rejected DATA-ONLY encounters can be re-approved in the future if enough data is later submitted to allow for a reasonable chance of identification of this animal. As part of the DATA-ONLY rejection process, an email is sent to the encounter submitter informing him/her of the DATA-ONLY status of the encounter and of the insufficiency of data needed for accurate identification. Some submitters may have additional photographs that can be added to each encounter to allow for approval and public display.
2. Click Permanently Delete to remove this encounter from the Library altogether. Note: Deleted encounters can be restored by a Library administrator. If you accidentally select Permanently Delete, email [email protected] and reference the encounter number that you deleted to have the encounter restored.
1.2.3 Reaccepting a DATA-ONLY encounter
If you subsequently receive enough data for a DATA-ONLY encounter to allow for possible reidentification in the future, you can reaccept the encounter, moving it out of a DATA-ONLY status.
To reaccept a DATA-ONLY encounter, click Reaccept in the blue bar at the left of a DATA- ONLY encounter (a.k.a. a rejected encounter) to move it back to an Unapproved state.
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2 Extracting Spot Patterns
The ECOCEAN Library is a mark-recapture framework for storing and analyzing whale shark data. Among its many features, it supports spot pattern searches to help identify new and previously sighted individual whale sharks using photographs and frame grabs from video.
2.1 Basic assumptions 1. For each encounter submitted to the Library, spot patterns can be added for the left
and/or right sides if one or both sides have been appropriately photographed. Ideal photographs are taken perpendicular to the spot pattern area, as defined and demonstrated in the red box below.
Figure 2. The area used for spot pattern extraction on the left and right sides is highlighted in red. This photo represents an
ideal photograph taken perpendicular to the spot pattern area.
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2. To prevent the double-counting of two separate encounters of the same shark as two separate animals (e.g. one encounter submitted with only a left side photo and a second encounter submitted with only a right side photo) only encounters with left side spot patterns that have been extracted and added to the Library can be allocated as new sharks. If they cannot be visually matched to another shark in the Library by other features(scars, etc.), encounters for which only right side spot data can be extracted, must remain“Unassigned” until they can be matched to another shark in the Library.
3. Once an encounter has been assigned to a shark, new spot data cannot be added to
it. This prevents the overwriting of existing spot data which was used to identify the encounter previously. To add spot data to an encounter that has been assigned to a shark, the encounter must first become “Unassigned” by removing it from the shark.
2.2 Pre-processing a photo for pattern recognition Any encounter that is not assigned to a shark can have left and/or right side spot data added to it if properly oriented photos were submitted for it. Here are the steps required to extract a pattern from a photo. Let’s use the following photo as a good example.
Figure 3. A new photograph submitted to the ECOCEAN Library.
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To ensure that a photograph is properly aligned for spot pattern recognition analysis using either the I3S or Modified Groth algorithms supported by ECOCEAN, it must first be pre-processed in a graphics package. There are a variety of free, inexpensive, and expensive software packages that you can use for the following steps. ECOCEAN uses Adobe Fireworks for simple photo processing or Paint.NET1, which is freely available. Other applications, such as Adobe Photoshop2, Adobe Fireworks3, and GIMP4 can also be used. The directions for using your choice of graphics packages will vary. The objective is to obtain a final image of the shark that is correctly-oriented as described in the following sections.
1 http://fileforum.betanews.com/detail/PaintNET/1096481993/1
2 http://www.adobe.com/products/photoship/index.html 3 http://www.adobe.com/products/fireworks/
4 http://www.gimp.org
2.2.1 Obtaining a source image from the ECOCEAN Library
1. Login to the Library through a web browser if you have not done so already.
2. In the Library, go to the encounter to which you want to add spot data.
3. In the encounter page, click on the photograph from which you want to extract spot data.
4. Click on the link “Click here to access the original source image.”
5. When the original image appears in your browser, right-click the image and select Save Picture As to save the picture to your Desktop.
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Figure 4. Viewing an image and accessing the source file in the ECOCEAN Library.
2.2.2 Pre-processing an image in Paint.NET
If you choose to use Paint.NET to pre-process source images on Windows XP/Vista, use the following instructions. If you use another software program, replicate these basic steps using your tool.
1. Open Paint.NET on your computer.
2. Open the picture in Paint.NET by selecting Open from the File menu.
3. Use the Rectangle Select tool to select the spot pattern area used for whale sharks.
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Figure 5. A whale shark image loaded in Paint.NET.
4. From the Image menu, select Crop to Selection to reduce the image to only the needed patterning area.
Figure 6. Selecting the spot pattern area in Paint.NET.
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Figure 7. A cropped spot pattern area.
5. We also recommend reducing the size of very large images to ensure the spot pattern area fits neatly onto the Paint.NET canvas at 100% magnification. To adjust image
size, use the Rectangle Select tool to select the entire image and then select Resize from the Image menu. Adjust image size appropriately.
6. After obtaining a reduced image of only the spot pattern area, create a new layer to hold
a horizontal adjustment line using the Add New Layer button of the Layers pallet.
Figure 8. Adding a new layer to hold a reference line. 138
7. With the new layer selected in the Layers palette, add a horizontal reference line to the
image using the Line\Curve tool . Hold the Shift key when drawing the reference line to ensure it is perfectly horizontal.
Figure 9. Adding a horizontal reference line.
8. Select the Background layer containing the cropped spot patterning area and then use
the Rectangle Select tool to select the entire patterning area image. From the Layers menu, select Rotate/Zoom.
Figure 10. Accessing the Rotate tool.
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9. Use the Rotate and Pan controls of the Rotate/Zoom dialog box to rotate and move the spot patterning area until the vertebral column is flat against the horizontal line. Click OK when you are done.
Figure 11. Obtaining a proper orientation for the spot patterning area.
Note: The section of the vertebral column just above the fifth gill may curve downward slightly and not fit flush to the line. This is acceptable so long as the rest of the vertebral column above the pectoral fin is parallel to the line.
10. Select the layer with the horizontal red line in the Layers pallet and click the Delete Layer button to remove it.
11. Use the Rectangle Select tool and the Image, Crop to Selection menu command to reduce you image to only the needed spot patterning area.
12. Choose File, Save As to save your completed processed image under a new name.
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Figure 12. An image ready for spot pattern extraction.
You are now ready to extract the spots using ECOCEAN Interconnect and send them to the ECOCEAN Library.
2.3 Sending a pattern to the ECOCEAN Library with the ECOCEAN Interconnect Client The ECOCEAN Interconnect Client is a small standalone software application that analyzes the image you prepared above and sends spot data to the ECOCEAN Library where it can be used with the I3S and modified Groth algorithms to identify potential matches.
Note: To use Interconnect for the first time, you will need to download it and configure your computer to run it. See Client Software for information on how to obtain and install the free Interconnect client.
To extract a spot pattern and send it to the ECOCEAN Library:
1. Open the ECOCEAN Interconnect Client.
2. From the File menu, select Open left-side shark image. 141
Figure 13. Opening a new image in ECOCEAN Interconnect.
3. In the Open dialog box, select the pre-processed image file and then click Open.
Note: Only JPG and GIF images are usable at this time in Interconnect.
4. After the image appears in Interconnect, from the File menu select Spot Selection.
5. Select the three reference points needed for the I3S algorithm (the Modified Groth algorithm does not require these) by left-clicking on the appropriate locations in the image. The red text overlaying the image will prompt you. The order of spots is: i. Top of the 5th gill.
ii. Posterior point of the pectoral fin on the flank. If the fin is not horizontal,
select the point above it where the white countershading underneath meets the pigmented skin along the flank.
iii. Bottom of the 5th gill.
To unselect any reference point, right-click it.
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Figure 14. Selecting the three reference points needed for the I3S pattern recognition algorithm.
6. Left-click in the image to select the center points of all of the spots in region of interest.
To unselect any spot, right-click it.
Figure 15. Selecting spots for computer-assisted photo-identification.
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7. After you have selected all of your spots, from the Database menu, click Submit to ECOCEAN Library to send the spot data via the Internet to the ECOCEAN Library. You will need an open Internet connection to perform this step.
Note: Once an encounter is allocated to a shark in the Library, new spot data cannot be submitted for it unless it is removed from the shark to which it belongs. This limitation is designed to preserve spot data used to justify a match and to protect that spot data from accidental overwriting. If you are attempting to add spots to an encounter already assigned to a shark, you must first remove the encounter from the shark to add the spots.
8. In the Send a left\right- side pattern to the ECOCEAN Library dialog box, enter the encounter number in the ECOCEAN Library to assign the pattern to. Click OK when you are ready to send the pattern.
Figure 16. Entering the encounter number to assign the extracted spot pattern to.
9. A new browser window will open to confirm that your spots have been received and prompting you to send in your processed image generated in steps 1-16 above. You may also be prompted to login first.
Figure 17. Uploading the processed image.
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10. Browse, select, and submit the processed image in your browser.
11. Your image is now visible (when you are logged in) in the web page for the encounter.
12. Confirm that the appropriate spots have been extracted and mapped using the visual remapping capability in the Library. In the encounter’s web page in the Library, select the link Click here to see the spots mapped to the image, which appears above the image you submitted to see the spots remapped to the image and to confirm their accuracy.
Figure 18. The spot data file itself in the ECOCEAN Library.
Figure 19. Ensuring that submitted spots remap to the original image correctly in the ECOCEAN Library.
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3 Comparing Spot Patterns with sharkGrid
The sharkGrid allows connected computers to participate in spot pattern recognition scans run from the ECOCEAN Library. When you start a pattern matching scan for an encounter in the ECOCEAN Library, you are actually creating a scanTask made up of individual comparisons called scanWorkItems. The scanWorkItems of a scanTask are executed in efficient, parallel groups that are distributed across the grid of computers. The results are reassembled once all comparisons are complete. Each encounter can have up to two active scanTasks: one for a left side pattern and another for a right side pattern.
3.1 Starting a spot pattern comparison scan on sharkGrid Once you have extracted a spot pattern for an encounter, you can look for matches to it across a global set of patterns stored in the ECOCEAN Library. To start looking for matches:
1. Select a left-side or right-side radio button from the Find Pattern Match form in the Action/Edit bar.
2. Click Start Scan.
Figure 20. Staring a scan with the Find Pattern Match form
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Starting a pattern matching scan creates a new scanTask in sharkGrid. The scanTask is assigned a unique identifier defined as:
“scanL”+encounter number for a left-side scan
“scanR”+encounter number for right-side scan
Note: There can be only one left-side scanTask and one right-side scanTask for an encounter, whether they are completed or uncompleted. If you decide to redo a spot pattern and scan for a match again, you must first delete the old scanTask if it still exists.
You can manage your scanTasks in the sharkGrid Administration page by selecting sharkGrid from the Administer menu (login required).
3.2 Examining the results of a scanTask You can check the status of and view the results of scanTasks from the sharkGrid Administration page.
3.2.1 Pending scanTasks
Pending scanTasks are scanTasks that are:
• being created for submission to the grid
• being executed on the grid
• completed but whose results have not yet been written
Figure 21. Two pending scanTasks in sharkGrid waiting for results to be written
You can delete a pending scanTask by clicking the Delete button.
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You can examine the results of a pending scanTask (and move it to “Completed” status) by clicking Write Result. The scanTask will subsequently appear in the Completed scanTasks table.
3.2.2 Completed scanTasks
Completed scanTasks are scanTasks that have been successfully completed in sharkGrid and whose results have been written out for review.
Figure 22. Several completed scanTasks in sharkGrid waiting to be removed
You can delete a completed scanTask that you initiated by clicking the Delete button.
You can examine the results of a completed scanTask by clicking View.
Note: Please promptly delete any completed scanTasks after viewing the results. The latest results can always be reviewed from the relevant encounter page.
3.3 Administering sharkGrid This section describes how sharkGrid operates and how users with various levels of access can change its behaviour.
3.3.1 The sharkGrid queue: Adding and deleting scanTasks
The creation and deletion of scanTasks occurs in a single queue. This means that if several users have simultaneously created new scanTasks or attempted to delete existing ones, these changes will occur one by one. A new create/delete scanTask operation will only start after the previous one in the queue has finished. Therefore, you may not see the results of a
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scanTask delete (i.e. the removal of the scanTask from the sharkGrid Administration page) immediately.
The number of scanTask additions/deletions in the queue for Administrator-level users is visible in this section of the sharkGrid Administration page:
Figure 23. The scanTask queue in sharkGrid
3.3.2 Viewing connected client nodes and their performance
You can view information about connected client nodes doing work in sharkGrid from this section of the sharkGrid Administration page:
Figure 24. gridManager statistics presenting information about sharkGrid nodes and their efficiency
The following information is available for each
node:
• IP - the IP address of the nodeNodeID - a unique identifier for the node. This identifier is randomly generated.
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• #CPU - The number of CPUs available for processing on the node computer. sharkGrid will take advantage of multicore CPU architectures and run scanWorkItems in parallel on multicore machines.
• Targeted? - whether the client node in working on a specified scanTask or is generally available for processing all scanTasks.
• # Finished - the number of scanWorkItems (individual pattern comparisons) completed by the node.
• Chunk size - the number of scanWorkItems sent to the node after each request. All nodes start with one scanWorkItem initially and are given larger workloads with subsequent requests. Ultimately, chunk size is determined according to a pre-defined algorithm in the ECOCEAN Library.
Overall performance in sharkGrid is measured by:
• % inefficient collisions - this is the percentage of duplicate work performed by nodes in sharkGrid. The ideal is 0. A collision is detected when a node tries to check in a comparison already completed by another node.
• Total work items completed since startup - this is the number of total comparisons performed in sharkGrid since the last restart of the ECOCEAN Library.
3.3.3 Configuring sharkGrid behaviour
From the sharkGrid Administration page, you can configure the following performance parameters if you have Administrator privileges:
• Set number of allowed nodes - defines the maximum number of client nodes that can participate in sharkGrid
• Set node timeout - defines how many milliseconds can pass without a heartbeat from a client node before the node is considered to be no longer connected to sharkGrid
• Set checkout timeout - defines how long (in milliseconds) after a scanWorkItem is
checked out for processing that it can be checked out by another node. The assumption is that, after this timeout period, the node originally checking out the scanWorkItem has left the grid. A very small value for this may cause inefficient duplication of effort. A very large value may cause a slowdown in overall scanTask processing.
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• Set number of allowed scanTasks - this is the maximum number of simultaneous scanTasks allowed in sharkGrid. Once the maximum is reached, existing scanTasks must be deleted before new ones are added. Alternatively, the maximum could be increased.
• Set maximum chunk/group size sent to nodes - this is the maximum number of comparisons (scanWorkItems) sent to a node after a request. All nodes start at one and advance in group size with each request. Setting this value too high may cause nodes to run out of memory or to take too long processing and register inefficient collisions. A very small value causes nodes to spend more time requesting data than processing it, which is also inefficient.
4 Interpreting Pattern Match Results You can access the results of a pattern matching scan (a.k.a. scanTask) by:
• Clicking View for a completed scanTask in the sharkGrid Administration page
• Clicking Groth: Left/Right-side scan results or I3S: Left-side scan results from the related encounter page
The results are displayed in two tabs: Modified Groth and I3S.
Figure 25. Scan results
4.1 Modified Groth results Details for the inner workings of the Modified Groth algorithm are available in the paper:
Arzoumanian, Z., Holmberg, J. and Norman, B. (2005) An astronomical pattern-matching algorithm for computer-aided identification of whale sharks Rhincodon typus. Journal of Applied Ecology, 42 (6), 999-1011.
The Modified Groth results are displayed in two areas:
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4.1.1 Metric results
Metric results provide the quantitative and qualitative results of the algorithm.
Figure 26. Metric results for the Modified Groth algorithm
The metric results for each comparison are ranked from best potential match to least. The listed values are:
• Shark - The shark that the potentially matching pattern belongs to.
• Encounter - The encounter with the potentially matching pattern.
• Fraction matched triangles - The fraction of matched triangles between the two spot patterns. Possible values range from 0 to 1. The higher the value, the stronger the match.
• Match score - The final match score, as described in Arzoumanian et al. The higher the value, the stronger the match.
• logM std. dev. - The standard deviation of the logarithm of magnification differences between matched triangles in the two spot patterns. The lower this value, the stronger the match.
• Confidence - The qualitative confidence of the match.
• Matched keywords - The photo keywords that both potentially matched encounters share in common.
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4.1.2 Visual results
The visual results for the Modified Groth algorithm display matched spots between potentially matched patterns. To compare results, look for each colored spot on the left to match a corresponding spot in the pattern on the right. Click Next and Previous to tab through potential matches in order of match score.
Figure 27. Visual results for the Modified Groth algorithm
4.2 I3S results Details for the inner workings of the I3S algorithm are available in the paper:
Van Tienhoven, A.M., Den Hartog, J.E., Reijns, R.A., and Peddemors, V.M. (2007) A computer- aided program for pattern-matching natural marks on the spotted raggedtooth shark Carcharias taurus (Rafinesque, 1810). Journal of Applied Ecology (2007) 44, 273–280.
The I3S results are displayed in two areas:
4.2.1 Metric results
Metric results provide the quantitative and qualitative results of the algorithm.
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Figure 28. Numeric results of the I3S algorithm
The metric results for each comparison are ranked from best potential match to least. The listed values are:
• Shark - The shark that the potentially matching pattern belongs to.
• Encounter - The encounter with the potentially matching patten.
• Fraction matched triangles - The fraction of matched triangles between the two spot patterns. Possible values range from 0 to 1. The higher the value, the stronger the match.
• Match score - The final match score, as described in Van Tienhoven et al. The lower the value, the stronger the match. Values of “0.0” should be ignored.
4.2.2 Visual results
The visual results for the I3S algorithm display matched spots between potentially matched patterns. To compare results, look for each colored spot on the left to match a corresponding
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spot in the pattern on the right. Click Next and Previous to tab through potential matches in order of match score.
Figure 29. Visual results for the I3S algorithm
5 Matching Sharks This topic describes how to allocate matched and unmatched encounters as sharks in the ECOCEAN Library.
5.1 Creating a new shark If an approved encounter has a properly-oriented, left-side spot pattern extracted and added to the database, and if the spot pattern scan and visual inspection of the encounter photos do not produce a match to an existing shark, the encounter can be allocated as a new shark. This allocation is a first “mark” of the animal. Future sightings are considered “recaptures”.
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Figure 30: A perfectly-oriented left-side pattern required for new shark identification.
Does your left-side pattern look like Figure 30?
Notice that the top of the pattern area is perfectly horizontal, all spots in the pattern region are selected and the photographer was perpendicular to the patterning area. If your processed image does not look like this (or a slight variation of it), do not allocate it as a new shark. Creating new sharks with suboptimal extracted patterns lowers their potential for reidentification through pattern matching (Modified Groth and I3S) in the future.
To create a new shark from an encounter:
1. Login to the Library through a web browser if you have not done so already.
2. Go to the approved and unassigned encounter that you want to create a new shark from.
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3. Click Edit next to the Assigned to Shark field at the top of the encounter.
4. In the Create New Shark field that appears in the blue Action/Edit bar on the left, type a new name for the shark. The name of the shark should conform to the series code assigned to you in the Library. For example, sharks first identified at Ningaloo Reef are assigned names according to the format ‘A-XXX’. For example, ‘A-011’ is the eleventh shark identified in the Library at Ningaloo Reef.
5. Click Create. The Library will inform you of the success or failure of its efforts to allocate this encounter as a new shark. When you create a new shark, the submitter of the encounter is automatically informed via email of the updated status of their encounter as a newly identified shark in the Library.
Note: Once an encounter is allocated to a shark in the Library, new spot data cannot be submitted for it unless it is removed from the shark to which it belongs. This limitation is designed to preserve spot data used to justify a match and to protect that spot data from accidental overwriting.
5.2 Adding an encounter to an existing shark If a spot pattern scan or visual analysis shows that a new encounter is a match to a shark already identified in the Library, you can assign the encounter to the shark. This assignment represents a “recapture” of that animal. To assign an encounter to a shark:
1. Login to the ECOCEAN Library through a web browser if you have not done so already.
2. Go to the approved and unassigned encounter that you want to add to a shark.
3. Click Edit next to the Assigned to Shark field at the top of the encounter.
4. In the Add to Shark field that appears in the blue Action/Edit bar on the left, type the name of the existing shark to add the encounter to.
5. In the Matched By field, select:
• Pattern match ── for encounter matches made with the assistance of extracted spot patterns and database scans
• Visual inspection ── for encounter matches made from “by eye” visual comparison only
6. Click Add. A message confirming success or failure will appear in your browser. If successful, all submitters and photographers of the shark’s new encounter and its previous encounters are informed of the re-sight of this shark via email.
Note: Once an encounter is assigned to a shark in the Library, new spot data cannot be submitted for it unless it is removed from the shark to which it belongs. This limitation is
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designed to preserve spot data used to justify a match and to protect that spot data from accidental overwriting.
6 Searching Through the ECOCEAN Library The ECOCEAN Library offers the following capabilities to search through submitted data.
6.1 Encounter search The Encounter Search allows you to search through whale shark sightings reported to the ECOCEAN Library. You can filter the search according to:
• approved, rejected, or unapproved status of the encounter • sex • identification status and minimum number of sightings of the shark • length • location • location code • submitter or photographer name • date range
Results are also displayed in Google Maps for those reports where GPS coordinate were provided.
Figure 31. Google Maps result for an encounter search. 158
6.1.1 Data export
The Encounter Search allows filtered results to be exported in two formats:
• Excel with columns corresponding to Darwin Core values (Administrators only)
• Google Earth KML file (Google Earth version 5 or higher). Only encounters with reported GPS coordinates are added to the KML file.
1. Google Earth export options
The ECOCEAN Library provides one Google Earth export option: “Add a timestamp to the KML to animate on a timeline”. If the checkbox for this option is checked, the exported KML file will contain time data that allows you to play through sightings along a timeline in Google Earth 5+. The timeline starts at the date of the earliest sighting (with GPS coordinates) in the filtered results and ends with the date of the last encounter (with GPS coordinates).
Figure 32. Google Earth with timeline displayed for encounters with timestamps
2. Google Earth export icons
Three colors of icons are displayed in Google Earth results: blue for males, pink for females, and white for whale shark encounters of unknown sex. Some icons may also have a number in them. This number represents the shark length (rounded to the nearest whole meter) for the sighting, if reported.
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Figure 34. Google Earth icons for whale shark sightings interpreted as: male, female, unknown sex.
7 FAQs
1. Why do spots sometimes appear mismapped in the results of a scanTask?
Two things can happen when you see mismapped spots in a scan result.
a. The match results are stored in an XML file. However, the images they pull to map the matched spot pairs to are the latest extraction images. Therefore, when someone remapped the spots, your scan results did not change, but they're now mapped to the new extraction image that was uploaded. The best way to resolve this is:
1. Delete the old scanTask if it still exists.
2. Clear your browser cache. (The new Firefox 3 browser works well for this!)
3. Rerun the scanTask to get the new, corrected results.
b. Flash has a cache for images and patterns. This can also cause a spot mismap if you've looked at a scan result then remapped and rescanned. The second set of results might show the first mapping. In this case, clear your browser cache and then reload the page.
2. Why do some encounters show a "Bad File" thumbnail?
This situation can occur when the first submitted image/video is not of a supported image or video file type. This can be fixed by logging into the ECOCEAN Library and uploading a supported image/video type OR by asking the webmaster ([email protected] ) to resolve the bad file (if you don't have required access). If you upload a supported file type, follow the instructions below to regenerate the image thumbnail.
1. Log into the ECOCEAN Library.
2. Go to the appropriate encounter page.
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3. If you have edit permissions for the encounter, look in the Action/Edit bar for the Reset Thumbnail box. Choose the submitted image to generate the thumbnail from and then click Reset Thumbnail.
4. Go back to the page where you first encountered the “BAD FILE” thumbnail and Refresh/Reload the page in your browser. The updated thumbnail will appear.
3. How do you determine if a shark is "new"?
Looking at the process backwards, the end goal of whale shark mark-recapture is to obtain a large group of sharks with low misidentification for accurate population modeling. Our photo- identification standards, as presented in this wiki, are in place to help support accurate matching across the data sets of multiple users…the assumption being that some of us might be sharing sharks and therefore we need standards to ensure that we process data identically and can quickly and accurately match them between our data sets when they appear. The pattern recognition algorithms provide a reliable, fast way to do that. Ultimately we could get the same effect (very laboriously and less accurately) with “by eye” matching, but this adds the risk of double counting a shark if not properly matched to an existing photo across catalogues. The pattern recognition software we use (Modified Groth and I3S algorithms) significantly reduce this risk, and therefore we use them as a standard for all new sharks (i.e. the patterns provide a measure of statistical evidence that there is not a match elsewhere in the catalogues).
That said, there will always be some human-added variability and error, including some missing/extra mapped spots and of course variable angle between shark and photographer. We know that our probability of automated matching degrades (for both the I3S and Modified Groth algorithms) when the angle between photographer and shark flank falls away from perpendicular, spots outside the patterning area are added or too few spots from the patterning area are added.
The algorithms can internally account for some of this variability (Spot! helps too), but we have to make a judgment call for each unmatched pattern and peer review it. The questions we ask are:
• Are all of the proper spots mapped?
• Is the pattern properly rotated?
• Is the angle between photographer and shark appropriate, roughly within fifteen degrees of perpendicular on either side and with minimal roll?
Ultimately, it's a judgment call based on those criteria, and we have peer review and discussion (even sometimes disagreements) about whether a shark is new. We're simply trying to ensure that every new shark has the maximum probability of being matched in the future.
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Appendix 1A. How to Join sharkGrid
How to join sharkGrid : http://www.whaleshark.org/wiki/doku.php?id=how_to_join_sharkgrid
Welcome to the sharkGrid! The sharkGrid allows you to donate spare cycles of your computer to whale shark research. Specifically, the processing power of your computer can be “borrowed” for resource intensive tasks, such pattern recognition or data mining. sharkGrid uses “global volunteer computing” to distribute intensive tasks between multiple computers to allow them to complete much more quickly.
There are a few quick things you should know before participating in sharkGrid.
1. Once running, the sharkGrid client must stay open to allow your computer to aid in whale shark spot pattern processing. If for any reason you want to leave the grid, close the client. If you restart your computer, you must restart sharkGrid to participate.
2. Your Internet connection must remain open to participate in sharkGrid. sharkGrid obtains all patterns to compare from the World Wide Web.
3. Your computer must have a Java Runtime Environment (JRE) 5 or higher installed. You can download a JRE from the Sun Microsystems Java web site. The JRE is different from the Java Virtual Machine (JVM) that runs in your browser. The JRE allows Java applications to run as programs on your desktop.
4. Please disable any power saving features enabled on your computer. For example, many PCs will enter a suspended state (a.k.a. “System standby”) after a period of time with no user interaction. Other power saving schemes stop the hard drive(s), which will also interfere with the sharkGrid client.
5. sharkGrid participation is subject to the ECOCEAN Library Visitor Agreement. By visiting this web site and joining sharkGrid, you are agreeing to the terms and conditions therein.
6. sharkGrid can run in the background while you perform other tasks on your computer. However, sharkGrid is used for intensive computation, and other applications running on your computer simultaneously with sharkGrid may slow down.
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7. sharkGrid requires significant computing power. We recommend you only run sharkGrid on computers with 1 gigabyte (GB) of RAM or more.
1A.1 Downloading the sharkGrid client
You can download a zip file containing the sharkGrid client
(http://www.whaleshark.org/interconnect/sharkGridClient_1_2.zip).
1A.2 Installing and starting the sharkGrid client
To install the sharkGrid client to your desktop, follow these instructions.
Figure A.1. The sharkGrid client running successfully.
1A.2.1 Windows
1. Unzip the sharkGrid client zip file to a custom folder on your Desktop or elsewhere on your computer. This is the folder that sharkGrid will run from.
2. Determine how much memory is available on your computer.
3. In the sharkGrid folder, right-click StartGridClient.bat and select Edit. This is the command-line statement used to start sharkGrid:
java -classpath .;jdo2-api-2.0.jar;jpox-1.1.9.jar -Xms512m -Xmx1g workApplet3
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You should edit ONLY the parameter ”-Xmx1g” to match the memory on your computer. You can specify your memory as megabytes (m) or gigabytes (g). The following are sample settings:
”-Xmx1g” or ”-Xmx1024m” for 1 gigabyte of RAM, which is also 1024 megabytes.
”-Xmx2g” or ”-Xmx2048m” for 2 gigabytes of RAM, which is also 2048 megabytes.
4. Save your changes to StartGridClient.bat.
5. Double-click StartGridClient.bat. You should see the sharkGrid client start and begin looking for work to do.
6. If the sharkGrid client fails to start, you may need to manually set the JAVA_HOME environment variable on your computer. This variable tells your computer where Java is installed so that it can use it when the “java” command is issued to start the sharkGrid client. Follow these steps on Windows to set JAVA_HOME:
1. Right click on the My Computer icon on your desktop and select Properties.
2. Click the Advanced Tab.
3. Click the Environment Variables button.
4. Under System Variables, click New.
5. Enter the variable name as JAVA_HOME.
6. Enter the variable value as the base directory of the JRE, such as C:\Program Files\Java\jre1.5.0_09. The value you set depends on where the JRE was installed on your computer.
7. Click OK.
8. Double-click StartGridClient.bat. You should see the sharkGrid client start and begin looking for work to do.
1A.2.2 Linux
1. Unzip the sharkGrid client zip file to a custom folder on your Desktop or elsewhere on your computer. This is the folder that sharkGrid will run from.
2. Determine how much memory is available on your computer.
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3. This is the command-line statement used to start sharkGrid from the folder you unzipped it to:
java -classpath .:jdo2-api-2.0.jar:jpox-1.1.9.jar -Xms512m -Xmx1g workApplet3
You should edit ONLY the parameter ”-Xmx1g” to match the memory on your computer. You can specify your memory as megabytes (m) or gigabytes (g). The following are sample settings:
”-Xmx1g” or ”-Xmx1024m” for 1 gigabyte of RAM, which is also 1024 megabytes.
”-Xmx2g” or ”-Xmx2048m” for 2 gigabytes of RAM, which is also 2048 megabytes.
4. This command assumes that Java 5 or higher is already installed and available from the command line.
5. Create a shell script containing this command and consider creating a cron job to run this script during the times of the day that you know your machine will be idle.
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Appendix 1B. Photo Keywords Used in the ECOCEAN Library
The following images provide examples of the keywords used to tag photos in the ECOCEAN Library.
Figure B.1. Gill damage (left or right)
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Figure B.2. Horizontal pattern lines (left or right)
Figure B.3. Line doubled (left or right)
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Figure B.4. Nicks, fin, 1st dorsa
Figure B.5. Nicks, fin, 2nd dorsal
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Figure B.6. Nicks, fin, caudal lower
Figure B.7. Nicks, fin, caudal upper
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Figure B.8. Nicks, fin, pectoral left
Figure B.9. Propeller cuts, body
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Figure B.10. Propeller cuts, fin
Figure B.11. Scar, body (left or right)
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Figure B.12. Scar, bite
Figure B.13. Scar, head
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Figure B.14. Skin growth
Figure B.15. Tag
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Figure B.16. Truncation, fin, 1st dorsal
Figure B.17. Truncation, fin, caudal lower
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Figure B.18. Truncation, fin, caudal upper
Figure B.19. Truncation, fin, pectoral (left or right)
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Figure B.20. Twinned spots, fiducial (left or right)
This tag is used for twinned spots that fall within the patterning area used for computer-assisted photo-identification.
Figure B.21. Twinned spots, external (left or right)
This tag is used for twinned spots that fall outside the patterning area used for computer-assisted photo-identification.
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Appendix 1C. Spot!
The spot pattern recognition algorithms used in whale shark mark-recapture (Modified Groth and I3S) assume a flat, two-dimensional surface when analyzing the relationships between spots. While each algorithm has some tolerance for skew in an image, both quickly degrade in their ability to match identical patterns as the angles between those patterns increases.
Spot! allows you to map a skewed 2D image to a 3D whale shark model and obtain a properly- oriented left- or right-side pattern for use with the Modified Groth and I3S algorithms.
Figure C.1. Spot!
This perspective correction has been used in the ECOCEAN Library to match images taken from very extreme angles to previously tagged whale sharks. For example, Spot! was used to make this match for shark M-025 in the ECOCEAN Library.
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Figure C.2: Pattern match from Spot!
1C.1 Spot! requirements Spot! requires Java 5 or Java 6 on your computer. The Java Runtime Environment (JRE) can be downloaded from Sun Microsystems at: http://www.java.com
1C.2 Loading Spot! Spot! can be loaded from within the Client Software page of the ECOCEAN Library or directly from this link: http://www.whaleshark.org/spot/spot.jnlp
1C.3 Basic Spot! instructions
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There are just a few basic steps to using Spot!.
1. Load the skewed image into Spot! using the Open button.
2. Scale the image to the 3D model by holding down the middle mouse button and moving the mouse forward and backward.
3. Move the loaded image left and right by holding down the right mouse button.
4. Manipulate the 3D model by holding down the left mouse button on its edges and moving the mouse up, down, left, and right.
5. Repeat steps 2-4 to align the guidelines on the 3D to these features on the 2D image: 5th gill, lowest lateral line, and vertebral column.
6. Once aligned, click the Map button to map the 2D image to the 3D model.
7. Click the Export button to save the properly aligned image for spot mapping with ECOCEAN Interconnect
1C.4 Spot! FAQ 1. How accurate is Spot?
We're still testing it. So far, we've made a lot of very strong matches in our test cases using real- world data. Spot! has exceeded our expectations in its ability to match patterns from skewed images to properly-oriented patterns. As with any software tool, the result is as good as the user, and Spot! requires patient, careful alignment between 3D model and 2D image. At this time, we're using Spot! to go back through encounter reports and identify sharks from images that were previously unsuitable for existing pattern recognition algorithms. Statistical tests of Spot!'s accuracy will be performed in the near future. But again...it's always going to be only as good as its user.
2. Can I use Spot! to identify new whale sharks from skewed images?
That's a longer discussion about misidentification in mark-recapture population modeling. At this point, Spot! is NOT being used to identify new whale sharks in the ECOCEAN Library.
However, we may choose to do so in the future. We currently use it only to identify previously marked whale sharks. In general, we HIGHLY recommend the tandem use of both the Modified Groth and I3S pattern recognition algorithms in addition to rigorous peer review to reduce misidentification.
3. Can Spot! be used for other species?
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Yes! Spot! can load other 3D models (.obj file format) and map images to them. This allows researchers for other spotted species to use image catalogs and pattern recognition algorithms where the observer cannot orient the camera correctly to the animal, such as when camera traps are used. If you are interested in using Spot! and the dual pattern recognition algorithms of the ECOCEAN Library, please contact [email protected].
4. Is Spot! open source?
It will be as soon as our initial tests are complete. For now, it is freeware under license. Use it as you like.
5. Is a single whale shark model representative of the long-term morphology of an individual?
Probably not for very small (rare) and very large whale sharks, such as those found in the Galapagos Islands. The truncated 3D whale shark model used in Spot! is based on the proportions of average sized whale sharks, generally about 7 meters in total length (TL). We assume changes are proportional in all three dimensions for smaller and larger sharks (generally 5 m to 10 m in TL) within the range of most ecotourism activity. As our understanding of whale shark biology increases, we may use multiple models for different stages of growth.
6. Why is this model truncated at the front and back? Where are the fins?
The model focuses only on the fiducial region used for spot pattern recognition of whale sharks: left- and right-side patterning behind the gills. This area does not distort as much as many other regions of the shark during normal motion. By using a truncated model, we focus the eye on only the region of interest and consider only the region of interest.
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Appendix 1D. TapirLink and the ECOCEAN Library
Certain data from the ECOCEAN Library are shared with other biology and ecology portals, such as the Global Biodiversity Information Facility and Fishbase. This sharing occurs at regular intervals via an installed TapirLink provider. Information shared fits the basic DarwinCore and Geospatial Extension specifications. The specifications map cleanly to the base “encounter” data structure displayed in the ECOCEAN Library.
ECOCEAN, after consulting with some of its users, has adopted these restrictions on shared data:
• Publicly submitted data is shared by default
• Users with login access can manage which encounters are shared
• Photos are not shared due to copyright restrictions
• Spot pattern data is not shared to protect ongoing research initiatives
• Individual shark identifications are not shared to protect ongoing research initiatives
• Data is shared with the following restriction listed in the metadata: “Original photos and extracted spot patterns withheld. Please contact [email protected] regarding usage permissions.” Requests for usage of data submitted by users with ECOCEAN Library login access will be routed to the library user for the ultimate decision.
ECOCEAN as a general policy, but without strict enforcement, will not share viaTapirLink publicly submitted data for the current calendar year as a precaution to protect this vulnerable species. Individual users of the ECOCEAN Library retain the right to decide which encounters to share.
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Appendix 1E. ECOCEAN Library Location Codes
The following are the location codes used for standardized location identification in the ECOCEAN Library.
1a - Ningaloo, non-specific 1a1 - North Ningaloo, off Exmouth 1a2 - South Ningaloo, off Coral Bay 1b - Christmas Island 1c - Indonesia 1d - Malaysia 1e - Thailand 1f - Myanmar 2- Caribbean, non-specific 2a - Utila, Honduras 2b - Belize 2c - Mexico, East Coast 2d - Texas, Flower Gardens 2e - U.S. Gulf Coast 4a - Mozambique 4b - Tanzania 4c - Kenya 4d - South Africa permissions 4e- Djibouti 4f- Madagascar 5a - Maldives 5b - Seychelles 6a - Donsol, Philippines 6b - Taiwan 7 - eastern Pacific 7a - Galapagos Islands 7b - Cocos Island 7c - Panama, Pacific coast 8a - Great Barrier Reef
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Appendix 1F. ECOCEAN Whale Shark Photo-Identification Library Access Policy
This policy defines who may be given access to the ECOCEAN Library and how access may be requested.
Background
The ECOCEAN Whale Shark Photo-identification Library (“ECOCEAN Library”) is privately funded by ECOCEAN, a non-profit organization in Australia. The ECOCEAN Library provides standardized research software and analytical techniques for the study of whale sharks (Rhincodon typus). Access to the ECOCEAN Library is provided free of charge to individuals or organizations selected according to the criteria of this policy. The number of new accounts that can be provided for the ECOCEAN Library each year is limited by:
• ECOCEAN's available resources to support existing users
• ECOCEAN's available resources for existing whale shark research efforts
Requesting access
Requests for access to the ECOCEAN Library should be sent via email to: [email protected]. In the request, please state:
• Your research objectives • The organization you represent (if appropriate) • The timeframe of your research • How ECOCEAN Library tools and data could benefit your research • How much data you have already collected for your research • Your qualifications
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Request review
Requests for access to the ECOCEAN Library are reviewed and voted upon by the Management Committee of ECOCEAN within 60 days. The Management Committee may also request outside assistance in reviewing your application.
The Management Committee weighs these and other criteria when reviewing a request for access:
• Your ability and willingness to contribute new whale shark data
• Your ability and willingness to follow the analytical techniques used in the ECOCEAN Library
• Your ability and willingness to develop new techniques for whale shark research
• Your ability and willingness to develop new science for other fields using whale shark data
• Your ability and willingness to use whale shark data collaboratively
• ECOCEAN's available resources to properly support you
The final decision for a request is made by a majority decision of the ECOCEAN Management Committee. Responses are provided via email. The Management Committee wishes to acknowledge in advance that not all valid requests for access can be approved due to limitations on available resources.
By invitation
The Management Committee of ECOCEAN may choose to invite appropriate individuals to participate in the ECOCEAN Library without a formal request for access. Invitations for access require a majority decision by the Management Committee and are reviewed in accordance with the criteria defined above.
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Appendix 1G. ECOCEAN Library Visitor Agreement
Welcome to the ECOCEAN Library! Please read this Visitor Agreement. By using this web site, you accept its terms. This Visitor Agreement applies to any web page using the following domains, which are collectively known as the “ECOCEAN Whale Shark Photo-identification Library” or in shortened form “ECOCEAN Library”:
whaleshark.org sharkgrid.org
The Internet is an evolving medium, and we may change the terms of this Visitor Agreement from time to time. By continuing to use any of the ECOCEAN Library sites after we post any such changes, you accept the Visitor Agreement, as modified. We may change, restrict access to, suspend or discontinue the ECOCEAN Library, or any portion of the ECOCEAN Library, at any time.
If you disagree with any material you find in the ECOCEAN Library, we suggest that you respond by noting your disagreement in an email to [email protected]. We invite you to bring to our attention any material you believe to be factually inaccurate. Please forward a copy of the material to our webmaster along with an explanation of your disagreement.
If you are an owner of intellectual property who believes your intellectual property has been improperly posted or distributed via the ECOCEAN Library, please notify us immediately by sending email to our webmaster.
A link to another Web site does not constitute an endorsement of that site (nor of any product, service or other material offered on that site) by the ECOCEAN Library or its participants.
NO SOLICITING
You agree not to use the ECOCEAN Library to advertise or to solicit anyone to buy or sell products or services, or to make donations of any kind, without our express written approval.
USE OF MATERIALS
Any photographs that you submit to the ECOCEAN Library remain YOUR intellectual property, and the ECOCEAN Library and its participants agree not to use them for media purposes without your express permission. However, by submitting photographs and whale shark sighting data you give ECOCEAN and its participants permission to use this data for research and conservation purposes. Data, such as shark identifications, may be derived from your submissions. This data becomes the intellectual property of the ECOCEAN Library and may not be published or re-used without the express permission of the ECOCEAN Library.
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The Internet allows people throughout the world to share valuable information, ideas and creative works. To ensure continued open access to such materials, we all need to protect the rights of those who share their creations with us. Although we make the ECOCEAN Library freely accessible, we don't intend to give up our rights, or anyone else's rights, to the materials appearing in the ECOCEAN Library. The materials available through the ECOCEAN Library are the property of the ECOCEAN Library or, in the case of photographs and images, the property of individual contributors. All photographs and data are protected by copyright, trademark and other intellectual property laws. You may not reproduce any of the materials without the prior written consent of the owner. You may not distribute copies of materials found on the ECOCEAN Library in any form (including by email or other electronic means), without prior written permission from the ECOCEAN Library.
Requests for permission to use, reproduce, or distribute materials found in the ECOCEAN Library should first be sent to webmaster at whaleshark dot org. Requests will be evaluated and responded to (yes or no) as quickly as possible. Our main concern is to protect intellectual property and to ensure that credit is given where credit is due. Our mission is to facilitate global cooperation within the whale shark research community, and we are working to make as much data as possible available while protecting the rights of individual contributors.
LINKING
We welcome links to the ECOCEAN Library. You are usually free to establish a hypertext link to any of the ECOCEAN Library pages so long as the link does not state or imply any sponsorship of your site by the ECOCEAN Library. Pages linking to the Library should include, to the best of your ability, factually correct information about the ECOCEAN Library and about whale sharks. In other words, please respect the scientific mission of the ECOCEAN Library and help us ensure that only accurate information about whale sharks is disseminated.
FRAMING
No Framing. Without the prior written permission of the ECOCEAN Library, you may not frame any of the content in the ECOCEAN Library, or incorporate into another Web site or other service any intellectual property of the ECOCEAN Library or its data contributors. Requests for permission to frame our content may be sent to: webmaster at whaleshark dot org.
DISCLAIMER OF WARRANTIES AND LIABILITY
We work hard to make the ECOCEAN Library interesting and informative, but we cannot guarantee that our users will always find everything to their liking. Please read this Disclaimer carefully before using the ECOCEAN Library.
YOU AGREE THAT YOUR USE OF THE ECOCEAN LIBRARY IS AT YOUR SOLE RISK. BECAUSE OF THE NUMBER OF POSSIBLE SOURCES OF INFORMATION AVAILABLE THROUGHOUT, AND THE INHERENT HAZARDS AND
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UNCERTAINTIES OF ELECTRONIC DISTRIBUTION, THERE MAY BE DELAYS, OMISSIONS, INACCURACIES OR OTHER PROBLEMS WITH SUCH INFORMATION. IF YOU RELY ON ANY ECOCEAN LIBRARY MATERIAL, YOU DO SO AT YOUR OWN RISK. YOU UNDERSTAND THAT YOU ARE SOLELY RESPONSIBLE FOR ANY DAMAGE TO YOUR COMPUTER SYSTEM OR LOSS OF DATA THAT RESULTS FROM ANY MATERIAL AND/OR DATA DOWNLOADED FROM OR OTHERWISE PROVIDED THROUGH THE ECOCEAN LIBRARY. THE ECOCEAN LIBRARY IS PROVIDED TO YOU AS IS, WITH ALL FAULTS, AND AS AVAILABLE. UNDER NO CIRCUMSTANCES SHALL THE PARTICIPANTS, PROGRAMMERS, AND CONSULTANTS IN THE ECOCEAN LIBRARY BE LIABLE TO YOU OR ANYONE ELSE FOR ANY DAMAGES ARISING OUT OF USE OF THE ECOCEAN LIBRARY, INCLUDING, WITHOUT LIMITATION, LIABILITY FOR CONSEQUENTIAL, SPECIAL, INCIDENTAL, INDIRECT OR SIMILAR DAMAGES, EVEN IF WE ARE ADVISED BEFOREHAND OF THE POSSIBILITY OF SUCH DAMAGES. (BECAUSE SOME STATES DO NOT ALLOW THE EXCLUSION OR LIMITATION OF CERTAIN CATEGORIES OF DAMAGES, THE ABOVE LIMITATION MAY NOT APPLY TO YOU. IN SUCH STATES, THE LIABILITY OF ECOCEAN AND ITS STAFF AND AFFILIATES IS LIMITED TO THE FULLEST EXTENT PERMITTED BY SUCH STATE LAW.)
USER ACCOUNTS
The ECOCEAN Library staff does its best to ensure that information we post to the ECOCEAN Library is timely, accurate, and scientifically valuable. To obtain access to certain services of the ECOCEAN Library, you may be given an opportunity to register with the ECOCEAN Library. As part of any such registration process, you will be provided an user name and a password. You agree that the information you supply during that registration process will be accurate and complete and that you will not register under the name of, or attempt to enter the ECOCEAN Library under the name of, another person. You will be responsible for preserving the confidentiality of your password, sharing it with no one else without express permission, and will notify the staff of the ECOCEAN Library of any known or suspected unauthorized use of your account. You agree to indemnify, defend and hold harmless ECOCEAN, its affiliates and participants, and their officers, directors, employees, agents, licensors and suppliers, from and against any and all losses, expenses, damages and costs (including reasonable attorneys' fees) resulting from any violation of this Visitor Agreement or any activity related to your account (including negligent or wrongful conduct) by you or any other person accessing the ECOCEAN Library using your account.
MISCELLANEOUS
In the event that any portion of this Visitor Agreement is found to be invalid or unenforceable for any reason, such invalidity or unenforceability shall not affect the enforceability or validity of any other portion of this Visitor Agreement, which shall remain in full force and effect and be construed as if the invalid or unenforceable portion were not part of the Visitor Agreement.
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By using the ECOCEAN Library, you agree to abide by the terms of this Visitor Agreement. We hope you enjoy using the ECOCEAN Library, and we welcome suggestions for improvements.
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Appendix 1H. User Agreement ‘ECOCEAN Library’
The following policy applies only to individuals with login access to the ECOCEAN Library.
ECOCEAN Whale Shark Photo-identification Library Usage Agreement Subscribers: Please read the following Usage Agreement logging in to the ECOCEAN Whale Shark Photo-identification Library. By logging into the ECOCEAN Whale Shark Photo- Identification Library, you agree to all of the terms and conditions of this ECOCEAN Whale Shark Photo-identification Library Usage Agreement (“Usage Agreement”), including the terms, conditions and notices contained in the “Usage” section of this Usage Agreement. If you do not agree with ANY of the terms or conditions contained herein, please do not use the ECOCEAN Whale Shark Photo-identification Library. Please contact webmaster @whaleshark. org for any questions related to this agreement.
ECOCEAN reserves the right to change, modify, add or remove portions of this Usage Agreement or the terms or conditions contained herein. However, subscribers need to be notified 30 days prior to the changes taking place. If the Subscriber deems that they will no longer be able to meet their obligations under the User Agreement or that they will no longer be able to use or access the Service in a useful manner they must inform ECOCEAN directly and no longer use the ECOCEAN Whale Shark Photo-identification Library in any manner.
1. Definitions.
The ECOCEAN Whale Shark Photo-identification Library (the “Library”) is a suite of online informational services (the “Services”) provided by ECOCEAN, consisting of software applications and content provided by members of ECOCEAN, members of the general public, and governmental management agencies. “You” or “yours” refers to each person or entity, as applicable, that subscribes to the ECOCEAN Whale Shark Photo-identification Library (the “Subscriber”).
Authorized Users
Authorized users are those persons, and only those persons, who have been issued a user identifier and password by ECOCEAN.
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2. General.
The User Account Request Form, this Usage Agreement and any other policies relating to the use of the ECOCEAN Whale Shark Photo-identification Library (collectively, this “Agreement”) set forth the terms and conditions that apply to your use of the ECOCEAN Whale Shark Photo-identification Library. By signing and submitting the User Account Request Form to ECOCEAN, you are deemed to have agreed to comply with all of the terms and conditions of this Agreement. The right to use the Services is limited to Subscribers and Authorized Users and is not transferable to any other person or entity. You are responsible for protecting the confidentiality of your access to the Services and for complying with any guidelines relating to security measures designed to prevent unauthorized access as outlined in the Usage Agreement.
You are responsible to make reasonable efforts to inform Authorized Users aware of the terms of use as outlined by this agreement. You are not liable for actions of other users but agree to work with ECOCEAN to rectify any problems caused by Authorized users who infringe upon the terms of the Agreement. If the Subscriber fails comply with any material term or condition of the Usage Agreement ECOCEAN, may terminate this Agreement upon written notice if the Subscriber does not cure such noncompliance within sixty (60) days of receiving written notice of the breach.
All Subscribers are authorized to provide remote access to Services only to Authorized Users as long as reasonable security procedures are undertaken that will prevent remote access by institutions or individuals that are not Authorized Users under this Usage Agreement.
3. Usage.
Your use of the Services constitutes your agreement to all of the terms, conditions and notices below in addition to the general terms and conditions contained in this Agreement. If you do not agree with these provisions, please do not use the Services and request an immediate termination of your account.
In General
As a condition of using the Services you agree to abide by all applicable local, Provincial, national and international laws and regulations relevant to the use of the Services including, without limitation, any applicable privacy legislation and policies. In addition, you warrant that you will not use the Services for any purpose that is unlawful or prohibited by this Agreement (including, without limitation, any use that infringes another’s copyright rights). You may not use the Services in any manner that could damage, disable, overburden or impair the Web site or any user of the Web site, or interfere with any other party’s use of the Services. You will not use the Services or content of the Library to accumulate data for or
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promote a Whale Shark Photo- identification Library that may in any way be deemed by ECOCEAN to be competitive with its own efforts.
Good Faith Data Collection, Reporting, Sharing, and Collaboration
As a Subscriber, you have the right to submit photographs, data, and content to the Library. The Library is a community resource, and its content is used by a number of different individuals and agencies for a variety of research and conservation purposes. In all cases, you will submit content and encounter reports as completely and as accurately as possible, obtaining permission to use any copyrighted materials before submitting them to the Library. In addition, you will not submit any content that uses language or imagery (verbal or visual) that is deemed as offensive by anyone for any reason. ECOCEAN reserves the right to edit your content to enforce this.
While you retain the copyright to your photographic data, you are agreeing to share this information with other Subscribers and to a limited extent with the general public. No requests for content “hiding” from other Subscribers will be honoured by ECOCEAN, though our security system does prevent content “tampering” and protects your critical data. Additionally, you agree not to compete with users of the ECOCEAN Library from other regions or to allow others to compete through your access. The ECOCEAN Library is for collaboration, and users must respect the individual research interests of others outside their region of interest.
Publications Using Data from the Library
In any case where the content or Services of the Library are used in any way to contribute to any publication (online or print), Subscribers must make a good faith effort to include a visible acknowledgment of the ECOCEAN Library in their publications. Furthermore, any Subscriber taking copyrighted content directly from the Library and using it in any publication or medium must first obtain the written consent of the appropriate copyright holder(s) (such as encounter photographers) and comply with all national and international copyright laws.
Copyright and Trademark Protection
All materials contained on the Services (including, without limitation, the Web site’s “look and feel,” layout, data, design, text, software, images, graphics, video and audio content (the “Content”) are the property of ECOCEAN or the individual contributors of the content (“the Owner”), and their rights are protected by copyright, trademark and other intellectual property laws and international treaties. You may not reproduce any of these materials without the prior written consent of the owner. You may not distribute copies of materials found on this web site in any form (including by email or other electronic means) without prior written permission from the owner.
Downloading Materials
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You may not publish, copy, automatically browse or download, display, distribute, post, transmit, perform, modify, create derivative works from or sell any Materials that you did not personally submit, information, products or services obtained from the Services in any form or by any means, including, without limitation, electronic, mechanical, photocopying, recording or otherwise, except as expressly permitted under applicable law or as described in this Usage Agreement. You also may not engage in systematic retrieval of data or other content or Materials from the Services to create or compile, directly or indirectly, a collection, compilation, database or directory. Nor may you “mirror” on your own site or any other server any Material contained on the Services, including, without limitation, the Services’ home page or result pages without the express and written consent of ECOCEAN. Use of the content and Materials on the Services for any purpose not expressly permitted by this Usage Agreement is prohibited.
Third Party Web Sites
Hyperlinks to other Internet resources are provided for your convenience. ECOCEAN has selected these resources as having some value and pertinence, but such resources’ development and maintenance are not under the direction of ECOCEAN. Thus, the content, accuracy, opinions expressed and other links provided by these resources are neither verified by ECOCEAN editors nor endorsed by ECOCEAN. Because ECOCEAN has no control over such Web sites and resources, you acknowledge and agree that ECOCEAN is not responsible for the availability of such external Web sites or resources. In addition, you acknowledge and agree that ECOCEAN does not endorse and is not responsible or liable for any content, advertising, products or other materials on or available from such Web sites or resources. Furthermore, you acknowledge and agree that ECOCEAN will not be liable, directly or indirectly, for any damage or loss caused by the use of any such content, products or materials.
4. Intellectual Property Rights.
You acknowledge that the Services contain copyrighted material, trademarks, and other proprietary information owned by ECOCEAN and individual contributors to the Library, and that your subscription does not confer on you any right, title or interest in or to the Services, the related documentation or the intellectual property rights relating thereto other than the rights you retain on the material that you directly submitted to the Library. Unauthorized copying of any portion of the Services may constitute an infringement of applicable copyright, trademark or other intellectual property laws or international treaties and may result in litigation under applicable copyright, trademark or other intellectual property laws or international treaties and loss of privileges granted pursuant to this Agreement.
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5. Account and Security.
You are responsible for maintaining the confidentiality of your method of accessing the Services.
6. Disclaimer of Warranty; Limitation of Liability.
YOU EXPRESSLY AGREE THAT USE OF THE SERVICES IS AT YOUR SOLE RISK. NEITHER ECOCEAN, ITS AFFILIATES NOR ANY OF THEIR RESPECTIVE EMPLOYEES, AGENTS, THIRD PARTY CONTENT PROVIDERS OR LICENSORS WARRANT THAT THE SERVICES WILL BE AVAILABLE AT ANY PARTICULAR TIME, UNINTERRUPTED, OR ERROR FREE; NOR DO THEY MAKE ANY WARRANTY AS TO THE RESULTS THAT MAY BE OBTAINED FROM USE OF THE SERVICES, OR AS TO THE ACCURACY, RELIABILITY OR CONTENT OF ANY INFORMATION OR SERVICE PROVIDED THROUGH THE SERVCIES. THE SERVICESARE PROVIDED ON AN “AS IS” BASIS WITHOUT WARRANTIES OF ANY KIND, EITHER EXPRESS OR IMPLIED, INCLUDING BUT NOT LIMITED TO WARRANTIES OF TITLE OR IMPLIED WARRANTIES OF MERCHANTABILITY OR FITNESS FOR A PARTICULAR PURPOSE, OTHER THAN THOSE WARRANTIES WHICH ARE IMPLIED BY AND INCAPABLE OF EXCLUSION, RESTRICTION OR MODIFICATION UNDER APPLICABLE LAW. IN NO EVENT SHALL ECOCEAN BE LIABLE TO YOU OR ANY OTHER PERSON FOR LOSS OF BUSINESS OR PROFITS, OR FOR ANY INDIRECT, INCIDENTAL OR CONSEQUENTIAL DAMAGES ARISING OUT OF THE USE OF, OR INABILITY TO USE, THE SERVICES, EVEN IF ECOCEAN WAS PREVIOUSLY ADVISED OF THE POSSIBILITY OF SUCH DAMAGES, OR FOR ANY OTHER CLAIM BY A SUBSCRIBER, AUTHORIZED USER, OR ANY OTHER PERSON. THIS WARRANTY GIVES YOU SPECIFIC LEGAL RIGHTS, AND YOU MAY ALSO HAVE OTHER RIGHTS WHICH VARY BY LOCATION.
In the event any claim relating to the performance or nonperformance by ECOCEAN pursuant to this Agreement, or in any other way concerning the Services, is made by a Subscriber or Authorized User, the actual damages to which such Subscriber or Authorized User may be entitled shall be limited to the lesser of the fees paid by the Subscriber or Authorized User for the Services or One US Dollar (US $1).
7. Indemnification.
To the maximum extent permitted by law, you agree to defend, indemnify and hold harmless ECOCEAN, its affiliates and their respective directors, officers, employees and agents from and against any and all claims and expenses, including attorneys' fees, arising out of the use or unauthorized copying of the Services or any of their content, the violation
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of this Agreement or any applicable laws or regulations, or arising out of your violation of any rights of a user.
8. Term and Termination of Agreement.
Either party shall have the right to terminate this Agreement at any time by providing notice of termination to the other party in accordance with the Subscription Form. In the event of termination of this Agreement by either party, you shall have no claims against ECOCEAN, its affiliates, or any individual contributors to the Library. Termination of this Agreement automatically terminates your license to use the Services, any content or any other materials contained therein.
9. Miscellaneous.
This Agreement is entire and complete, and no representations, warranties, agreements or covenants, express or implied, of any kind or character whatsoever have been made by either party hereto to the other, except as expressly set forth in this Agreement. Except as provided herein, this Agreement may not be modified or changed unless the same shall be in writing and signed by an authorized officer of the party to be bound thereby.
You may not assign any of your rights or delegate any of your obligations under this Agreement without ECOCEAN's prior written consent. If any provision of this Agreement is held to be overly broad in scope or duration by a court of competent jurisdiction such provision shall be deemed modified to the broadest extent permitted under applicable law. If any provision of this Agreement shall be held to be illegal, invalid or unenforceable by a court of competent jurisdiction, the validity, legality and enforceability of the remaining provisions shall not, in any way, be affected or impaired thereby. No waiver by either party of any breach or default hereunder shall be deemed to be a waiver of any preceding or subsequent breach or default. The section headings used herein are for convenience only and shall not be given any legal import.
The provisions of Sections 4, 6, 7 and 8 shall survive termination of this Agreement.
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Appendix 2
Individual whale shark recorded from multiple international sighting locations.
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Appendix 2: Individual whale shark recorded from multiple international sighting locations.
Locations where sighted n Approximate minimum straight line distance (km)
Identified Shark #
Date of first sighting
Date of last sighting
Maximum number of years between sightings
Belize, Honduras 13 250 H-001 H-006 H-008 H-015 H-016 H-017 H-046 H-051 BZ-011 BZ-014 BZ-016 BZ-019 BZ-021
03.04.1999 08.05.2001 05.04.1999 01.04.2005 13.04.2005 30.04.2002 17.04.2007 01.01.2002 05.04.1999 22.03.2003 03.06.2007 26.05.2008 27.04.2000
03.05.2007 06.05.2014 30.06.2006 09.06.2009 27.01.2012 02.05.2005 02.06.2010 23.04.2005 13.04.2009 13.04.2012 11.05.2012 02.03.2010 10.04.2012
8 13 7 4 7 3 3 3 10 9 5 2 12
Belize, Mexico (Atlantic) 9 250 BZ-002 BZ-008 BZ-007 BZ-009 BZ-012 BZ-023 MXA-008 MXA-740 MXA-959
06.05.2002 01.04.1999 23.04.2003 01.04.2002 23.04.2003 26.05.2008 12.06.2004 04.08.2005 03.08.2007
19.06.2013 14.07.2011 13.08.2012 09.08.2011 27.07.2011 30.07.2013 17.06.2013 24.05.2012 25.05.2013
11 12 9 9 8 5 9 7 6
Honduras, Mexico (Atlantic) 23 500 H-079 H-014 H-019 H-025 H-027 H-028 H-031
18.06.2009 24.04.2004 24.04.2005 10.12.2005 14.02.2005 13.02.2005 27.04.2005
19.08.2012 17.08.2012 31.07.2013 14.08.2011 23.07.2012 16.05.2011 19.06.2013
3 8 8 6 7 6 8
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H-032 H-041 H-048 H-049 H-054 H-057 H-058 H-066 H-071 H-081 H-087 H-090 MXA-049 MXA-437 MXA-577 MXA-718
26.10.2006 06.03.2006 23.03.2007 14.04.2007 10.09.2005 15.03.2008 06.02.2008 10.02.2005 21.06.2005 15.04.2010 20.09.2010 26.10.2006 15.02.2005 03.09.2009 01.03.2004 05.08.2006
19.08.2013 16.01.2013 26.07.2013 08.08.2013 02.01.2008 10.08.2013 04.09.2012 26.06.2013 30.01.2009 13.09.2012 28.07.2013 03.10.2013 22.07.2013 07.08.2013 22.08.2010 23.08.2010
7 7 6 6 3 5 4 8 4 2 3 7 8 4 6 4
Belize, Honduras, Mexico (Atlantic) 7 500 BZ-001 BZ-026 H-030 H-035 H-052 MXA-008
07.08.2002 27.04.2000 01.10.2005 01.01.1999 04.06.2007 12.06.2004
26.07.2013 30.09.2013 14.08.2011 29.07.2013 01.09.2012 13.08.2012
11 13 6 14 5 8
Belize, Honduras, Mexico (Atlantic), USA
1 1300 H-021 24.04.2000 10.07.2014 14
USA, Mexico (Atlantic) 9 800 GC-026 GC-047 GC-057 GC-058 MXA-030 MXA-255 MXA-291 MXA-343 MXA-970
22.06.2010 21.07.2006 11.09.2011 15.07.2009 06.07.2008 03.09.2009 23.12.2005 22.06.2010 11.06.2009
10.03.2012 22.06.2010 12.01.2013 15.09.2011 18.08.2011 05.10.2013 31.07.2013 04.09.2012 10.11.2010
2 4 2 2 3 4 8 2 1
USA, Honduras 1 1050 H-045 01.12.2002 22.08.2009 7
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Belize, USA, Mexico (Atlantic)
1 850 BZ-010 21.04.2003 19.06.2013 10
Mexico (Atlantic), Cuba
1 1000 MXA-301 12.06.2009 29.09.2013 4
Mozambique, South Africa 19 900 SA-002 SA-006 SA-007 SA-008 SA-010 SA-015 MZ-022 MZ-035 MZ-044 MZ-067 MZ-096 MZ-124 MZ-301 MZ-308 MZ-376 MZ-418 MZ-427 MZ-499 MZ-553
12.10.2006 14.10.2006 17.02.2007 12.05.2006 09.12.2008 05.04.2009 27.01.2007 19.04.2007 13.04.2007 18.07.2011 13.04.2007 17.12.2006 07.02.2007 01.06.2007 11.09.2009 14.01.2009 12.04.2009 08.12.2006 10.01.2007
18.01.2010 24.05.2009 09.06.2014 03.08.2012 05.03.2010 13.02.2010 24.03.2007 19.03.2012 24.01.2010 15.10.2013 09.12.2008 20.07.2013 19.05.2009 02.07.2011 24.09.2013 05.12.2011 23.10.2009 15.10.2009 21.08.2012
4 3 7 6 2 1 0 5 3 2 1 7 2 4 4 2 0 3 5
Mozambique, Tanzania 3 1500 MZ-029 MZ-129 MZ-136
14/04/2007 07.12.2006 21.11.2006
12.01.2014 13.12.2013 10.11.2012
7 7 6
Philippines (Donsol, Leyte) 2 500 P-220 P-237
23.02.2009 02.04.2009
04.06.2013 26.04.2013
4 4
Philippines (Donsol, Oslob) 2 500 P-259 P-448
23.04.2009 17.03.2010
13.01.2013 12.06.2012
4 2
Philippines (Oslob, Leyte) 6 300 P-391 P-429 P-456 P-464 P-555 P-556
15.01.2011 20.04.2012 30.05.2012 14.12.2011 10.04.2013 12.04.2013
02.05.2013 18.08.2012 12.04.2013 04.04.2013 21.05.2013 21.05.2013
2 0 1 2 0 0
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Oman, Qatar 5 500 OM-006 OM-030 OM-045 OM-046 Q-048
04.07.2009 18.09.2010 21.10.2011 14.10.2011 09.07.2011
20.09.2012 01.06.2012 01.06.2012 18.07.2012 12.07.2012
3 2 1 1 1
UAE, Oman 2 350 UAE-002 UAE-007
31.07.2009 14.04.2010
09.04.2010 15.04.2011
1 1
Costa Rica, Panama 1 200 CR-012 28.01.2010 05.01.2011 1
Bahamas, Dominican Republic, British West Indies (Turks and Caicos)
1 800 CRB-008 02.01.2013 10.02.2013 0
Malaysia, Thailand 1 800 T-049 12.07.2009 11.11.2009 0
Philippines (Leyte), Taiwan
1 1600 P-545 31.05.2012 06.04.2013 1
Australia, Indonesia 1 2700 A-424 01.07.2007 16.04.2012 5
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Appendix 3a-c
Acoustic Tag deployments at Ningaloo Marine Park (2011, 2012, 2013).
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Appendix 3a: Acoustic Tag deployments at Ningaloo Marine Park (2011).
Acoustic Tag No.
Shark No.
Date tagged Photo-ID Detections post tag-deployment in 2011
Acoustic Receiver detections in 2011
Maximum number of days between detections
Acoustic recording on day of tagging
64091 A-101 09.09.11 09.09.11
14.09.11, 15.09.11 17.09.11, 18.09.11 13.10.11, 16.10.11 24.10.11
35 NO
64093 A-579 09.09.11 09.09.11
16.09.11, 17.09.11 21.09.11, 11.10.11 17.10.11, 18.10.11 29.10.11, 31.10.11 02.11.11, 03.11.11 04.11.11, 05.11.11 06.11.11, 09.11.11 10.11.11, 12.11.11 13.11.11, 14.11.11 17.11.11, 18.11.11 20.11.11, 21.11.11 22.11.11, 26.11.11 29.11.11, 02.12.11 11.12.11, 12.12.11
20 NO
64092 A-108 11.09.11 11.09.11
11.09.11, 12.09.11 14.09.11, 19.09.11 18.10.11, 19.10.11 22.10.11, 31.10.11 06.11.11, 22.11.11 23.11.11, 25.11.11 27.11.11, 02.12.11
33 YES
64094 *A-411 13.09.11 13.09.11
14.9.11, 16.09.11 03.10.11, 4.10.11 28.10.11
95 NO
64110 A-584 15.09.11 15.09.11 15.09.11, 16.09.11 09.10.11
25 YES
64112 A-029 20.09.11 20.09.11
20.09.11, 12.10.11 22 YES
* Shark A-411 was also recorded within NRETA on 02.01.12, 06.01.12, 07.01.12, 08.01.12, 09.01.12, 14.01.12, 16.01.12, 21.01.12, 22.01.12, 23.01.12, 09.02.12
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Appendix 3b: Acoustic Tag deployments at Ningaloo Marine Park (2012).
Acoustic Tag No.
Shark No.
Date tagged Photo-ID Detections post tag-deployment in 2012
Acoustic Receiver detections in 2012
Maximum number of days between detections
Acoustic recording on day of tagging
29121 A-546 21.05.12 21.05.12
29.05.12 18.06.12 05.07.12
25.05.12, 26.05.12 18.06.12
23 NO
29118 A-847 21.05.12 21.05.12 29.05.12 - NO
29119 A-848 22.05.12 22.05.12 22.05.12, 23.05.12 13.06.12 17.06.12 20.06.12
21 YES
29123 A-783 22.05.12 21.05.12 22.05.12 28.05.12
28.05.12, 29.05.12 30.05.12, 25.08.12 26.08.12, 27.08.12 09.09.12, 10.09.12 11.09.12, 29.09.12 30.09.12, 01.10.12
86 NO
29120 *A-815 22.05.12 22.05.12 27.05.12 01.10.12
23.05.12, 24.05.12 30.05.12, 31.05.12 07.06.12, 08.06.12 03.07.12, 09.07.12 11.07.12, 12.07.12 14.07.12, 15.07.12, 16.07.12, 01.09.12, 04.09.12, 05.09.12 30.09.12, 01.10.12 02.10.12, 22.10.12 06.11.12, 07.11.12 08.11.12, 16.11.12 30.11.12
50 NO
29122 A-273 22.05.12 22.05.12 28.05.12, 29.05.12 1 NO
28407 A-828 07.08.12 07.08.12 No detections - -
28409 A-844 09.09.12 09.09.12 11.09.12, 04.10.12 05.10.12, 15.10.12 18.10.12, 26.10.12
10 NO
* Shark A-815 was also recorded within NRETA on 18.01.13, 19.01.13, 04.02.13, 19.02.13, 20.02.13, 02.03.13, 05.03.13, 06.03.13, 04.03.13, 08.03.13, 09.03.13, 10.03.13, 11.03.13, 15.03.13, 16.03.13, 29.03.13, 04.04.13, 13.04.13, 16.04.13, 17.04.13, 24.04.13, 25.04.13
* Shark A-815 was also recorded via photo-identification on 16.05.13 with no acoustic tag attached
202
Appendix 3c: Acoustic Tag deployments at Ningaloo Marine Park (2013).
Acoustic Tag No.
Shark No.
Date tagged
Photo-ID Detections post tag-deployment in 2013
Acoustic Receiver detections in 2013
Date tag removed
Maximum number of days between detections
Acoustic recording on day of tagging
28405 A-546 09.04.13 09.04.13
04.06.13 16.04.13, 20.04.13 22.04.13, 23.04.13 26.04.13, 27.04.13 07.05.13, 08.05.13 09.05.13, 13.05.13 17.05.13, 18.05.13 19.05.13
19.05.13 0.14 NO
28405 A-579 15.06.13 15.06.13 29.06.13
15.06.13, 25.06.13 29.06.13, 30.06.13 06.07.13
0.40 YES
28404 A-087 05.05.13 05.05.13 05.05.13, 08.05.13 13.05.13, 16.05.13 18.05.13, 19.05.13 24.05.13
0.14 YES
28406 A-884 09.04.13 09.04.13 16.04.13, 18.04.13 20.04.13, 22.04.13 28.04.13, 01.05.13 12.05.13, 19.05.13 21.05.13, 25.05.13 25.05.13, 30.05.13 06.06.13, 01.07.13
0.07 NO
28408 A-236 10.04.13 10.04.13 21.04.13 23.04.13
10.04.13, 19.04.13 21.04.13, 22.04.13 23.04.13
12.05.13 0.60 YES
28408 A-795 06.06.13 06.06.13 07.06.13 08.06.13
07.06.13 09.06.13 3.000 NO
28408 A-749 19.06.13 19.06.13 27.07.13 09.08.13
25.06.13, 27.06.13 28.06.13, 04.07.13 11.07.13, 31.07.13 02.08.13, 22.08.13 24.08.13
0.33 NO
203
Appendix 4a-c
Residency Index for whale sharks sighted for two or more days from photo-identification data for the
2011, 2012, 2013 calendar years.
204
Appendix 4a: Residency Index for whale sharks sighted for two or more days from photo-identification data for the 2011 calendar year.
Marked Individual First seen Last seen Period # days seen Residency index
A-001 20.05.2011 3.06.2011 15 3 0.20
A-003 2.05.2011 23.06.2011 53 4 0.08
A-013 1.05.2011 31.05.2011 31 2 0.06
A-029 15.04.2011 20.09.2011 159 3 0.02
A-052 8.05.2011 11.05.2011 4 2 0.50
A-063 16.05.2011 13.07.2011 59 2 0.03
A-076 20.04.2011 24.04.2011 5 4 0.80
A-088 21.05.2011 12.06.2011 23 2 0.09
A-101 22.05.2011 9.09.2011 111 4 0.04
A-102 26.03.2011 8.07.2011 105 9 0.09
A-108 1.06.2011 11.09.2011 103 2 0.02
A-232 17.05.2011 23.05.2011 7 3 0.43
A-233 28.03.2011 9.04.2011 13 2 0.15
A-251 18.04.2011 11.05.2011 24 3 0.13
A-268 29.05.2011 29.07.2011 62 2 0.03
A-271 3.05.2011 6.08.2011 96 2 0.02
A-302 28.04.2011 3.05.2011 6 2 0.33
A-317 28.05.2011 28.06.2011 32 2 0.06
A-342 31.05.2011 11.08.2011 73 3 0.04
A-349 14.06.2011 21.06.2011 8 2 0.25
A-357 27.04.2011 29.05.2011 33 3 0.09
A-399 20.04.2011 25.04.2011 6 2 0.33
A-421 23.03.2011 26.05.2011 65 3 0.05
A-446 7.04.2011 9.04.2011 3 2 0.67
A-463 18.04.2011 30.05.2011 43 5 0.12
A-466 24.03.2011 3.05.2011 41 3 0.07
A-475 22.03.2011 23.04.2011 33 2 0.06
A-476 25.03.2011 26.04.2011 33 3 0.09
A-496 1.04.2011 15.05.2011 45 2 0.04
A-528 1.05.2011 7.05.2011 7 2 0.29
A-529 1.04.2011 19.07.2011 110 8 0.07
A-534 24.04.2011 3.09.2011 133 6 0.05
A-540 16.05.2011 9.07.2011 55 5 0.09
A-542 10.06.2011 28.07.2011 49 2 0.04
A-546 9.06.2011 11.06.2011 3 2 0.67
A-550 22.03.2011 7.06.2011 78 6 0.08
A-552 12.04.2011 17.06.2011 67 2 0.03
A-557 15.03.2011 17.05.2011 64 3 0.05
A-568 4.04.2011 3.05.2011 30 4 0.13
A-569 18.03.2011 25.03.2011 8 2 0.25
A-573 22.04.2011 5.05.2011 14 3 0.21
A-584 30.04.2011 15.09.2011 139 2 0.01
A-586 4.04.2011 20.06.2011 78 3 0.04
A-591 21.05.2011 29.05.2011 9 2 0.22
A-592 30.04.2011 12.07.2011 74 9 0.12
A-616 16.04.2011 12.05.2011 27 2 0.07
A-633 24.04.2011 8.05.2011 15 2 0.13
A-655 26.04.2011 29.04.2011 4 2 0.50
205
A-676 4.04.2011 6.04.2011 3 2 0.67
A-682 2.04.2011 17.06.2011 77 5 0.06
A-703 17.05.2011 5.06.2011 20 2 0.10
A-705 1.05.2011 17.06.2011 48 3 0.06
A-707 8.06.2011 12.06.2011 5 3 0.60
A-712 30.03.2011 4.04.2011 6 2 0.33
A-726 30.04.2011 2.05.2011 3 2 0.67
A-728 16.05.2011 18.05.2011 3 2 0.67
A-737 26.04.2011 2.05.2011 7 2 0.29
A-1060 14.04.2011 29.04.2011 16 2 0.13
Mean 41.84 2.98 0.20
SE 5.24 0.22 0.03
* For the Residency Index calculation, those sharks where monitoring was only over two consecutive days were excluded
206
Appendix 4b: Residency Index for whale sharks sighted for two or more days from photo-identification data for the 2012 calendar year.
Marked Individual First seen Last seen Period # days seen Residency index
A-003 25.04.2012 1.07.2012 68 2 0.03
A-013 20.05.2012 2.07.2012 44 4 0.09
A-029 27.03.2012 29.04.2012 34 3 0.09
A-041 1.06.2012 9.09.2012 101 3 0.03
A-044 26.06.2012 29.06.2012 4 2 0.50
A-079 1.04.2012 28.05.2012 58 4 0.07
A-088 24.04.2012 16.06.2012 54 2 0.04
A-101 23.05.2012 25.05.2012 3 3 1.00
A-108 28.06.2012 27.08.2012 61 3 0.05
A-110 21.06.2012 9.07.2012 19 3 0.16
A-119 23.04.2012 14.05.2012 22 3 0.14
A-125 26.05.2012 21.07.2012 57 12 0.21
A-136 23.03.2012 22.06.2012 92 3 0.03
A-141 11.04.2012 22.06.2012 73 2 0.03
A-143 24.04.2012 17.06.2012 55 2 0.04
A-163 17.04.2012 28.06.2012 73 3 0.04
A-174 23.05.2012 10.07.2012 49 4 0.08
A-184 23.05.2012 29.06.2012 38 4 0.11
A-225 26.05.2012 23.06.2012 29 6 0.21
A-232 8.06.2012 23.06.2012 16 4 0.25
A-236 18.06.2012 5.07.2012 18 3 0.17
A-247 1.06.2012 6.07.2012 36 3 0.08
A-251 20.06.2012 3.07.2012 14 2 0.14
A-266 12.07.2012 19.07.2012 8 3 0.38
A-267 25.05.2012 2.06.2012 9 2 0.22
A-268 1.05.2012 17.05.2012 17 4 0.24
A-271 2.06.2012 19.07.2012 48 8 0.17
A-302 11.05.2012 29.06.2012 50 3 0.06
A-328 17.04.2012 16.07.2012 91 5 0.05
A-333 16.04.2012 16.07.2012 92 4 0.04
A-342 12.06.2012 19.07.2012 38 3 0.08
A-348 30.04.2012 3.06.2012 35 5 0.14
A-357 29.04.2012 4.07.2012 67 7 0.10
A-387 23.04.2012 17.06.2012 56 3 0.05
A-391 9.07.2012 15.07.2012 7 3 0.43
A-393 14.05.2012 5.06.2012 23 3 0.13
A-399 1.04.2012 1.07.2012 92 9 0.10
A-404 28.03.2012 7.07.2012 102 7 0.07
A-411 15.03.2012 28.04.2012 45 3 0.07
A-418 27.03.2012 23.07.2012 119 6 0.05
A-424 8.04.2012 18.05.2012 41 3 0.07
A-425 6.04.2012 1.06.2012 57 7 0.12
A-440 25.04.2012 20.06.2012 57 8 0.14
A-446 29.04.2012 16.07.2012 79 7 0.09
A-453 7.04.2012 12.07.2012 97 3 0.03
A-455 2.05.2012 4.05.2012 3 2 0.67
A-462 30.03.2012 25.06.2012 88 8 0.09
A-471 17.04.2012 2.06.2012 47 4 0.09
207
A-472 16.05.2012 18.05.2012 3 2 0.67
A-475 19.05.2012 22.06.2012 35 3 0.09
A-477 2.04.2012 30.06.2012 90 9 0.10
A-482 27.05.2012 11.06.2012 16 4 0.25
A-484 9.04.2012 1.06.2012 54 3 0.06
A-485 30.04.2012 8.05.2012 9 3 0.33
A-493 10.05.2012 2.07.2012 54 8 0.15
A-500 17.04.2012 7.07.2012 82 5 0.06
A-516 17.04.2012 19.05.2012 33 2 0.06
A-519 24.03.2012 4.07.2012 103 3 0.03
A-528 1.07.2012 9.07.2012 9 2 0.22
A-533 20.05.2012 6.07.2012 48 2 0.04
A-535 16.05.2012 27.05.2012 12 3 0.25
A-540 14.05.2012 1.07.2012 49 6 0.12
A-545 19.05.2012 13.07.2012 56 2 0.04
A-546 21.05.2012 5.07.2012 46 4 0.09
A-569 18.04.2012 16.06.2012 60 4 0.07
A-576 16.04.2012 8.05.2012 23 3 0.13
A-579 26.06.2012 22.07.2012 27 4 0.15
A-580 27.05.2012 3.07.2012 38 4 0.11
A-583 27.04.2012 3.07.2012 68 6 0.09
A-585 6.04.2012 19.05.2012 44 10 0.23
A-588 11.05.2012 29.06.2012 50 5 0.10
A-590 21.04.2012 4.08.2012 106 11 0.10
A-592 21.03.2012 11.07.2012 113 4 0.04
A-593 24.04.2012 6.05.2012 13 2 0.15
A-603 15.04.2012 27.06.2012 74 4 0.05
A-609 23.04.2012 5.07.2012 74 3 0.04
A-610 27.03.2012 7.07.2012 103 7 0.07
A-616 31.03.2012 7.06.2012 69 5 0.07
A-619 14.04.2012 17.05.2012 34 2 0.06
A-621 28.03.2012 30.06.2012 95 9 0.09
A-625 28.06.2012 4.07.2012 7 2 0.29
A-635 2.05.2012 6.07.2012 66 5 0.08
A-643 13.06.2012 3.07.2012 21 3 0.14
A-648 26.03.2012 13.07.2012 110 8 0.07
A-655 29.05.2012 9.07.2012 42 2 0.05
A-658 4.05.2012 11.06.2012 39 2 0.05
A-661 22.04.2012 21.05.2012 30 2 0.07
A-666 4.04.2012 13.07.2012 101 18 0.18
A-668 6.04.2012 10.07.2012 96 5 0.05
A-670 15.06.2012 15.07.2012 31 7 0.23
A-673 17.05.2012 26.05.2012 10 3 0.30
A-676 17.04.2012 1.09.2012 138 4 0.03
A-682 10.04.2012 11.05.2012 32 5 0.16
A-686 14.06.2012 1.07.2012 18 3 0.17
A-702 28.04.2012 4.07.2012 68 6 0.09
A-704 3.07.2012 6.07.2012 4 2 0.50
A-709 11.05.2012 2.07.2012 53 2 0.04
A-720 16.04.2012 18.05.2012 33 3 0.09
A-742 7.04.2012 18.07.2012 103 5 0.05
A-743 14.04.2012 4.07.2012 82 4 0.05
208
A-745 14.04.2012 17.04.2012 4 2 0.50
A-746 17.04.2012 23.06.2012 68 5 0.07
A-747 17.04.2012 18.05.2012 32 3 0.09
A-749 30.05.2012 18.06.2012 20 3 0.15
A-753 11.04.2012 5.07.2012 86 3 0.03
A-758 2.04.2012 21.06.2012 81 2 0.02
A-762 11.05.2012 14.07.2012 65 4 0.06
A-766 2.05.2012 23.05.2012 22 3 0.14
A-769 17.04.2012 21.04.2012 5 2 0.40
A-771 19.04.2012 23.06.2012 66 2 0.03
A-773 6.05.2012 4.06.2012 30 2 0.07
A-777 19.05.2012 13.06.2012 26 5 0.19
A-781 10.05.2012 19.06.2012 41 2 0.05
A-783 26.04.2012 28.05.2012 33 4 0.12
A-784 22.05.2012 2.06.2012 12 2 0.17
A-787 14.05.2012 18.05.2012 5 3 0.60
A-791 13.05.2012 22.06.2012 41 3 0.07
A-793 19.06.2012 14.07.2012 26 4 0.15
A-798 25.05.2012 30.05.2012 6 2 0.33
A-800 31.05.2012 7.06.2012 8 2 0.25
A-804 11.06.2012 27.06.2012 17 4 0.24
A-811 13.04.2012 6.07.2012 85 3 0.04
A-812 25.05.2012 5.07.2012 42 2 0.05
A-815 22.05.2012 1.10.2012 133 3 0.02
A-817 17.05.2012 9.06.2012 24 2 0.08
A-818 14.05.2012 1.07.2012 49 5 0.10
A-825 19.06.2012 23.06.2012 5 2 0.40
A-829 26.06.2012 14.07.2012 19 3 0.16
A-832 26.06.2012 6.07.2012 11 2 0.18
A-833 27.04.2012 27.06.2012 62 4 0.06
A-834 11.05.2012 3.07.2012 54 3 0.06
A-836 23.06.2012 26.06.2012 4 2 0.50
A-837 7.05.2012 30.05.2012 24 2 0.08
A-840 31.03.2012 9.07.2012 101 4 0.04
A-842 25.06.2012 11.07.2012 17 2 0.12
A-844 31.05.2012 9.09.2012 102 2 0.02
A-849 1.07.2012 18.07.2012 18 6 0.33
A-853 17.05.2012 30.05.2012 14 2 0.14
A-856 5.06.2012 19.06.2012 15 3 0.20
A-872 19.06.2012 22.06.2012 4 2 0.50
A-874 8.07.2012 12.07.2012 5 2 0.40
A-919 25.05.2012 5.07.2012 42 2 0.05
A-953 31.05.2012 1.07.2012 32 2 0.06
Mean 46.97 3.90 0.15
SE 2.74 0.20 0.01
* For the Residency Index calculation, those sharks where monitoring was only over two consecutive days were excluded
209
Appendix 4c: Residency Index for whale sharks sighted for two or more days from photo-identification data for the 2013 calendar year.
Marked Individual First seen Last seen Period # days seen Residency index
A-013 23.03.2013 25.05.2013 64 7 0.11
A-041 26.04.2013 26.05.2013 31 4 0.13
A-044 2.04.2013 18.06.2013 78 2 0.03
A-079 16.04.2013 23.04.2013 8 2 0.25
A-101 25.03.2013 11.09.2013 171 5 0.03
A-108 13.07.2013 26.07.2013 14 2 0.14
A-125 17.03.2013 23.07.2013 129 5 0.04
A-136 1.04.2013 15.05.2013 45 2 0.04
A-141 2.05.2013 26.05.2013 25 2 0.08
A-225 13.05.2013 2.07.2013 51 2 0.04
A-232 16.06.2013 14.07.2013 29 2 0.07
A-233 3.07.2013 5.07.2013 3 2 0.67
A-236 9.04.2013 23.04.2013 15 4 0.27
A-247 6.04.2013 13.06.2013 69 2 0.03
A-268 1.04.2013 7.11.2013 221 5 0.02
A-302 25.05.2013 29.06.2013 36 2 0.06
A-333 4.06.2013 3.07.2013 30 3 0.10
A-334 2.07.2013 11.07.2013 10 2 0.20
A-348 22.05.2013 15.06.2013 25 3 0.12
A-356 18.06.2013 23.07.2013 36 2 0.06
A-357 14.05.2013 18.05.2013 5 2 0.40
A-380 6.07.2013 13.07.2013 8 2 0.25
A-387 3.05.2013 30.07.2013 89 8 0.09
A-399 15.04.2013 1.06.2013 48 3 0.06
A-404 5.05.2013 30.05.2013 26 2 0.08
A-411 23.04.2013 26.04.2013 4 3 0.75
A-425 13.04.2013 31.08.2013 141 3 0.02
A-453 1.04.2013 9.05.2013 39 2 0.05
A-471 20.05.2013 7.07.2013 49 2 0.04
A-477 4.04.2013 12.07.2013 100 4 0.04
A-481 24.05.2013 7.07.2013 45 4 0.09
A-493 23.05.2013 8.07.2013 47 5 0.11
A-496 4.04.2013 26.06.2013 84 4 0.05
A-516 5.04.2013 11.07.2013 98 5 0.05
A-519 7.04.2013 21.07.2013 106 4 0.04
A-521 3.06.2013 29.06.2013 27 2 0.07
A-534 16.06.2013 12.07.2013 27 2 0.07
A-540 7.05.2013 2.07.2013 57 2 0.04
A-542 13.04.2013 17.08.2013 127 8 0.06
A-545 14.03.2013 3.09.2013 174 4 0.02
A-546 9.04.2013 4.06.2013 57 2 0.04
A-557 21.05.2013 25.06.2013 36 3 0.08
A-569 5.05.2013 29.08.2013 117 2 0.02
A-576 8.06.2013 2.07.2013 25 2 0.08
A-579 18.05.2013 29.06.2013 43 3 0.07
A-580 3.06.2013 8.08.2013 67 2 0.03
A-583 24.04.2013 26.04.2013 3 2 0.67
A-585 17.04.2013 9.07.2013 84 4 0.05
210
A-588 18.05.2013 18.07.2013 62 2 0.03
A-590 16.04.2013 5.07.2013 81 2 0.02
A-593 7.04.2013 25.04.2013 19 3 0.16
A-616 27.04.2013 12.06.2013 47 2 0.04
A-633 14.08.2013 24.08.2013 11 2 0.18
A-635 25.03.2013 15.04.2013 22 3 0.14
A-657 5.04.2013 30.05.2013 56 4 0.07
A-666 22.06.2013 7.08.2013 47 4 0.09
A-670 28.03.2013 9.04.2013 13 2 0.15
A-676 27.04.2013 12.08.2013 108 7 0.06
A-686 29.03.2013 13.09.2013 169 3 0.02
A-702 18.04.2013 18.07.2013 92 5 0.05
A-720 23.07.2013 21.08.2013 30 2 0.07
A-728 6.06.2013 21.08.2013 77 3 0.04
A-730 31.03.2013 9.04.2013 10 3 0.30
A-734 27.05.2013 1.09.2013 98 2 0.02
A-737 11.04.2013 19.04.2013 9 3 0.33
A-742 3.04.2013 22.07.2013 111 6 0.05
A-745 29.04.2013 12.05.2013 14 2 0.14
A-749 19.06.2013 9.08.2013 52 3 0.06
A-766 12.04.2013 1.06.2013 51 3 0.06
A-781 27.04.2013 9.07.2013 74 2 0.03
A-788 25.05.2013 7.07.2013 44 3 0.07
A-792 25.05.2013 10.07.2013 47 2 0.04
A-793 18.03.2013 21.08.2013 157 4 0.03
A-795 14.05.2013 8.06.2013 26 5 0.19
A-796 22.05.2013 1.07.2013 41 2 0.05
A-799 6.04.2013 18.05.2013 43 2 0.05
A-814 5.07.2013 9.07.2013 5 2 0.40
A-816 10.04.2013 2.06.2013 54 3 0.06
A-818 17.04.2013 2.09.2013 139 4 0.03
A-822 28.06.2013 23.07.2013 26 3 0.12
A-829 13.06.2013 4.07.2013 22 3 0.14
A-840 2.04.2013 8.09.2013 160 4 0.03
A-841 12.04.2013 11.07.2013 91 3 0.03
A-844 6.04.2013 10.04.2013 5 3 0.60
A-847 28.06.2013 3.07.2013 6 2 0.33
A-868 30.06.2013 18.07.2013 19 2 0.11
A-872 25.03.2013 3.07.2013 101 2 0.02
A-883 10.04.2013 25.04.2013 16 2 0.13
A-884 19.03.2013 9.04.2013 22 2 0.09
A-885 18.04.2013 22.06.2013 66 8 0.12
A-888 31.03.2013 4.05.2013 35 2 0.06
A-892 29.04.2013 4.05.2013 6 2 0.33
A-894 18.06.2013 4.07.2013 17 3 0.18
A-895 6.07.2013 19.07.2013 14 2 0.14
A-901 8.06.2013 16.07.2013 39 3 0.08
A-906 21.04.2013 19.06.2013 60 3 0.05
A-907 11.07.2013 29.09.2013 81 3 0.04
A-909 17.05.2013 2.07.2013 47 2 0.04
A-910 7.05.2013 28.05.2013 22 2 0.09
A-914 7.03.2013 31.08.2013 178 7 0.04
211
A-915 29.04.2013 4.07.2013 67 4 0.06
A-916 26.06.2013 3.08.2013 39 6 0.15
A-928 2.04.2013 16.07.2013 106 2 0.02
A-931 6.04.2013 24.04.2013 19 2 0.11
A-936 15.04.2013 18.04.2013 4 2 0.50
A-938 30.04.2013 21.05.2013 22 3 0.14
A-940 20.04.2013 12.06.2013 54 3 0.06
A-956 29.05.2013 23.08.2013 87 3 0.03
A-964 29.04.2013 22.06.2013 55 2 0.04
A-973 23.03.2013 6.04.2013 15 2 0.13
A-974 5.04.2013 18.07.2013 105 3 0.03
A-979 16.04.2013 9.06.2013 55 3 0.05
A-1000 28.05.2013 10.09.2013 106 3 0.03
A-1005 3.07.2013 14.07.2013 12 2 0.17
A-1035 1.04.2013 2.05.2013 32 3 0.09
A-1042 1.04.2013 12.05.2013 42 2 0.05
A-1044 21.05.2013 4.07.2013 45 2 0.04
A-1068 17.05.2013 17.07.2013 62 2 0.03
A-1074 30.06.2013 2.07.2013 3 2 0.67
A-1089 8.06.2013 20.07.2013 43 3 0.07
A-1111 22.05.2013 4.07.2013 44 2 0.05
A-1135 20.06.2013 11.09.2013 84 2 0.02
Mean 55.46 3.01 0.12
SE 4.07 0.13 0.01
* For the Residency Index calculation, those sharks where monitoring was only over two consecutive
212
