Journal of Fish Biology (2009) 74, 706–714
doi:10.1111/j.1095-8649.2008.02155.x, available online at http://www.blackwell-synergy.com
Deep-diving behaviour of a whale shark Rhincodon typus
during long-distance movement in the western
Indian Ocean
J. M. B RUNNSCHWEILER *†, H. B AENSCH ‡, S. J. P IERCE §
D. W. S IMS k{
AND
*Institute of Zoology, University of Zurich, Winterthurerstrasse 190, CH–8057 Zurich,
Switzerland, ‡The Open University, Walton Hall, Milton Keynes MK7 6AA, U.K., §Manta
Ray & Whale Shark Research Centre, Tofo Beach, Mozambique, kMarine Biological
Association of the United Kingdom, The Laboratory, Citadel Hill, Plymouth PL1 2PB,
U.K. and {Marine Biology and Ecology Research Centre, School of Biological Sciences,
University of Plymouth, Drake Circus, Plymouth PL4 8AA, U.K.
(Received 30 May 2008, Accepted 8 November 2008)
A whale shark Rhincodon typus satellite tagged off the coast of Mozambique showed a highly
directional movement across the Mozambique Channel and around the southern tip of
Madagascar, a minimum distance of 1200 km in 87 days. Dives to depths well into the
mesopelagic and bathypelagic zones (1286 m maximum depth) were recorded in a bathymetrically non-constraining habitat. The water temperature range recorded during the fish’s
# 2009 The Authors
movement was 34–299° C.
Journal compilation # 2009 The Fisheries Society of the British Isles
Key words: deep diving; dive behaviour; foraging; migration; ocean stratification; satellite
tagging.
Whale sharks Rhincodon typus Smith play an important role in socio-economic
activities in many parts of the world because they are both harvested by fisheries and the subject of major marine tourism industries (Davis et al., 1997;
Clarke et al., 2005; Stewart & Wilson, 2005). Although detailed knowledge
of the biology and ecology of R. typus remains limited, the discovery of several
predictable aggregations in highly productive areas has enabled much to be
learnt since Colman (1997) (Martin, 2007; Stevens, 2007). In terms of their
movements, R. typus satellite tagged in the Sea of Cortez, in South-east Asia
and off Western Australia, seemingly travel distances of thousands of kilometres, moving through multiple political jurisdictions (Eckert & Stewart,
2001; Eckert et al., 2002; Wilson et al., 2006). A detailed understanding of
†Author to whom correspondence should be addressed at present address. Swiss Federal Institute of
Technology, Raemistrasse 101, CH-8092 Zurich, Switzerland. Tel.: þ41 44 632 2071; fax: þ41 44 362 5085;
email: [email protected]
706
2009 The Authors
Journal compilation # 2009 The Fisheries Society of the British Isles
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RHINCODON TYPUS CROSSING THE MOZAMBIQUE CHANNEL
707
the vertical and horizontal movement patterns contributes to a better understanding of the still little-known habitat of this planktivorous species, and
which is important for sustainable management of socio-economic activities.
Despite an increasing amount of data on large-scale movement patterns in
R. typus, information on habitat use away from shallow-water feeding grounds
is generally lacking, especially in poorly studied, remote geographic regions.
The Indian Ocean is one of the most productive areas for R. typus sightings
and the species occurs all along the east African coast from the Red Sea to
South Africa (Compagno, 2001). A focal point for R. typus aggregation is
the coastline around Tofo in southern Mozambique (Cliff et al., 2007). Here,
a high density of fish gathers year-round in a narrow (c. 20 km2) corridor close
to shore and the high sighting rates and accessibility of the fish has led to the
development of a burgeoning marine tourism industry. Although the broaderscale movement patterns and behaviour of these fish are unknown, the local
population structure (81% males) suggests that these fish constitute a sub-set
of a larger population (unpubl. data). Satellite tags were deployed from this site
to gain a better understanding of both regional population linkages and also
the vertical habitat use of this planktivorous species.
On 18 February 2006, a male (shark 1; tag 1) and female (shark 2; tag 2)
R. typus were equipped with pop-up satellite archival tags (PTT-100; Microwave
Telemetry, Inc.; www.microwave-telemetry.com) at 24106° S; 35504° E, southeast of Tofo, Mozambique. The sizes of the fish were visually estimated to be 8
m (shark 1) and 6–7 m (shark 2), respectively, using the length of the research
vessel as reference. The fish were approached by a snorkeller, and the tags were
embedded into the dorsal musculature between the first and the second dorsal
fins using a spear gun. The full set-up consisted of the pop-up satellite archival
tag unit, a monofilament line (c. 100 mm) marked with an individually coloured plastic tube and a double barbed stainless steel anchor (modified Floy
FH-69 dart; Floy Tag & Mfg Inc.; www.floytag.com). The tags archived ambient light measurements (every 2 min), temperature and pressure readings (every
15 min) and recorded time of sunrise and sunset for subsequent calculation of
latitude and longitude. Error ranges for the temperature and pressure sensors
were 02° C and 27 m, respectively. In both tags, a constant pressure
release mechanism was enabled. This release feature is standard on Microwave
Telemetry pop-up satellite archival tags and automatically releases the tag and
begins to transmit to the Argos satellite if it senses that it has been at a constant
depth for 4 days. Depth variations <20 m were regarded as constant depth by
the tag model used in this study. The tags were programmed to pop-up on 30
November 2006. All times reported here (unless noted) are local times.
Both tags popped up prematurely after 7 (tag 1) and 87 (tag 2) days, respectively. Tag attachment duration was similar to other studies that satellitetagged R. typus using similar attachment methods (Graham et al., 2006; Wilson
et al., 2006). Tag 1 popped up on 25 February 2006 before 0900 hours at
24214° S; 35431° E (location class 3 on 26 February at GMT 0045 hours)
due to activation of the constant pressure release mechanism. This location
was 14 km south-west of the tagging site. The light-based geolocation longitude
estimates indicate that the fish stayed within a 100 km area off coast for the
duration of this short track. Argos positions received and temperature readings
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708
J. M. BRUNNSCHWEILER ET AL.
recorded after pop-up on 25 February strongly indicated that the tag had
washed ashore. All (100%) archived depth and seawater temperature data
(n ¼ 653) were retrieved from tag 1. The maximum diving depth of shark
1 was 48 m and the male shark spent >90% of its time at depths between
0 and 20 m in isothermic water. The distribution of time at depth between
day and night was not different (w2 test, d.f. 4, P > 005). Mean S.D. and
median depths were 74 71 m and 54 m, respectively. Mean S.D. ambient
seawater temperature experienced was 272 15° C, median ¼ 277° C and
range ¼ 209–293° C.
Tag 2 popped up on 16 May 2006, 87 days after attachment. The tag
detached for unknown reasons (last depth reading was 1076 m) and floated
on the surface for 4 days (depth readings constantly 0 m) until it ‘uplinked’
to the Argos satellite receiving system. The first Argos location (24224° S;
47639° E; location class 2) was received on 20 May at GMT 1952 hours from
off the south-east coast of Madagascar [Fig. 1(a)]. Raw estimates of locations
between attachment and tag pop-up were calculated from recovered light-level
data by Microwave Telemetry, Inc. using a proprietary algorithm derived from
standard celestial algorithms (Wilson et al., 2007). A total of 387% of daily
latitude estimates and 75% of daily longitude estimates, respectively, were
available for analysis. Light-level longitude estimates are typically accurate
and robust (Teo et al., 2004), and therefore, only longitude values were used
to describe the horizontal west to east movement of shark 2.
The minimum point-to-point distance covered by the R. typus in 87 days was
c. 1200 km between Mozambique and the east coast of Madagascar. Combining light-level longitude estimates with archived depth readings indicate
45°E
40°E
50°E
ue
35°E
am
biq
0m
Mo
z
*
25°S
1000 m
Madagascar
2000 m
3000 m
4000 m
Madagascar
Basin
Mozambique
Basin
Depth (m)
(a)
25°S
5000 m
Madagascar Ridge
40°E
35°E
0
200
400
600
800
1000
1200
1400
55
50
45
40
35
30
17 February 27 February
45°E
50°E
Longitude° E
(b)
(c)
9 March
19 March
29 March
8 April
18 April
28 April
8 May
18 May
FIG. 1. (a) Bathymetric map of the geographic area where both Rhincodon typus (shark 1 and shark 2) were
satellite tagged ( ). The white cross on the south-east coast of Madagascar indicates the position
from where the first satellite uplink from tag 2 (shark 2) was received. (b) Time-depth series of the
entire track and (c) daily longitude estimates for shark 2.
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2009 The Fisheries Society of the British Isles, Journal of Fish Biology 2009, 74, 706–714
709
RHINCODON TYPUS CROSSING THE MOZAMBIQUE CHANNEL
a strongly directional movement by shark 2. Minimum mean S.D. movement
rate calculated as the minimum straight distance between two consecutive daily
longitude estimates was 13 13 km h1 (median ¼ 08 km h1; range ¼ 00–
61 km h1). These estimated mean minimum movement rates are well within
previously published estimates of daily movement rates in this species (Eckert &
Stewart, 2001; Wilson et al., 2006). The data showing the animal crossing the
Mozambique Channel from coastal Mozambique to south-eastern Madagascar
waters provides the first track of an individual connecting mainland Africa to
Malagasy waters, where occasional R. typus shark aggregations are also known
to occur (Jonahson & Harding, 2007).
Vertical movements and diel patterns in behaviour were analysed using
871% (n ¼ 7262) of the archived depth readings and 878% (n ¼ 7321) of
the archived seawater temperature readings available from tag 2. Mean S.D. depth and temperature during the entire track were 661 1184 m
(median ¼ 108 m; range ¼ 0–1286 m) and 239 43° C (median ¼ 251° C;
range ¼ 34–299° C), respectively. The fish showed a broad depth distribution,
spending 796% of the time shallower than 100 m [Fig. 2(a)]. Within the first
100 m of the water column, the majority of the fish’s time was spent close
to the surface, with 60% of the time spent between 0–10 m depths [Fig. 2(b)].
It spent 64% of its time in water >24° C [Fig. 2(c)]. No thermocline could be
detected, indicating the fish dived in well-mixed layers. This result agrees with
previous satellite tagging studies for this species (Graham et al., 2006; Wilson
et al., 2006).
Light-based geolocation longitude estimates from tag 2 indicate that the
female fish initially moved south along the Mozambique coast in shallow
coastal water reaching the minimum longitude of 34382° E on 7 March
(a)
(b)
0–100
200–300
300–400
400–500
>500
100
0–10
28–30
10–20
26–28
20–30
24–26
30–40
22–24
40–50
20–22
50–60
18–20
60–70
16–18
70–80
14–16
80–90
12–14
<12
90–100
50
0
50
100
Temperature (°C)
Depth (m)
100–200
(c)
100
50
0
50
100
40
20
0
20
40
Per cent time-at-depth
FIG. 2. Histograms of percentage time spent at depth for (a) 100 m intervals, (b) 0–100 m and (c)
percentage time spent in different water temperatures ( , day; , night) for Rhincodon typus 2.
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J. M. BRUNNSCHWEILER ET AL.
[Fig. 1(a), (b)]. Between the date of tagging and 7 March, the fish stayed in
shallow coastal habitats, spending 64% of its time in water <10 m deep, with
only occasional deeper dives to a maximum depth of 108 m [Table I and Fig. 1(b)].
The mean ambient seawater temperature experienced in coastal habitat was
275° C (Table I). Temperature records from both sharks 1 and 2 indicated that
the upper 0–40 m of the water column close to the coast was isothermal with
little vertical stratification. Shark 2 stayed significantly deeper during the night
while in shallow coastal waters (Mann–Whitney U-test, d.f. ¼ 1, P < 0001).
No difference was detected between night and day ambient seawater temperatures in this coastal habitat (Mann–Whitney U-test, d.f. ¼ 1, P > 005).
Compared to the coastal habitat, an opposite diel pattern was found in oceanic waters (Table I). While in the oceanic habitat, mean depths were significantly shallower at night than during the day (Mann–Whitney U-test, d.f. ¼
1, P < 0001), which was linked to the mean ambient seawater temperature
experienced being significantly warmer at night than during the day (Mann–
Whitney U-test, d.f. ¼ 1, P < 0001).
After moving into oceanic waters in the Mozambique Channel, the vertical
movements of the fish changed dramatically to regular epipelagic diving punctuated by periodic deep dives, experiencing a wide range of ambient water temperatures (Table I). The first dive of R. typus below 100 m was recorded on 6 March
and regular deep diving to well below 100 m was observed from 12 March
onwards [Fig. 1(b)]. After leaving shelf waters on 7 March, the fish entered
deeper coastal waters heading east towards the Mozambique Basin [Fig. 1(a)].
Crossing the deep waters (c. 5000 m maximum depth) of the southern Mozambique Channel into the Madagascar Basin was characterized by increased diving
activity compared to the shallow coastal habitat including dives to depths well
into the mesopelagic and bathypelagic zones [Fig. 1(b)]. On 20 March, one dive
profile included descents to 1264 and 1092 m, respectively. Seawater temperatures recorded at these depths were 42 and 92° C, respectively. A third deep
dive to 1087 m with ambient seawater temperature of 55° C was recorded on
23 March. Between the beginning of April and 19 April, the R. typus crossed
the Madagascar Ridge entering the Madagascar Basin [Fig. 1(a)]. On 18 April,
the fish dived to at least 1286 m. Previous studies have found that R. typus dive
TABLE I. Summary statistics for dive data from shark 2 in coastal and oceanic habitats.
All depth and temperature variables show range (median) and mean S.D.
Coastal
Depth
Readings
Overall (m)
Day (m)
Night (m)
Temperature
Readings
Overall (° C)
Day (° C)
Night (° C)
Journal compilation
Oceanic
1395
0–108 (54) 82 129
0–54 (0) 61 85
0–108 (54) 103 158
1469
183–299 (278) 275 224–299 (278) 276 183–297 (278) 274 #
15
13
17
5867
0–1286 (215) 799 1277
0–1087 (323) 1182 147
0–1286 (108) 415 899
5852
34–295 (244) 23 43
52–295 (23) 215 5
34–29 (248) 245 26
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RHINCODON TYPUS CROSSING THE MOZAMBIQUE CHANNEL
711
to at least 980 m (Graham et al., 2006; Wilson et al., 2006). The maximum depth
of 1286 m, some 22 m deeper than the maximum diving depth recorded for
a basking shark Cetorhinus maximus (Gunnerus) in the North Atlantic Ocean
(Gore et al., 2008), shark 2 obtained in this study is, as far as is known, the
deepest dive depth ever directly recorded for any elasmobranch species. Deeper
dives are possible but cannot be recorded by this tag type since the maximum
possible recordable depth is 1286 m, triggering an immediate tag release and
pop-up if this depth is maintained for >15 min (Microwave Telemetry, pers.
comm.). The seawater temperature of 34° C recorded at this depth was the lowest temperature measured during the entire track (Table I). Bathypelagic dive
profiles were similar and showed the fish diving straight down from the epipelagic zone to the deepest recorded depth of each individual profile. This was followed by an ascent through the water column by undertaking very regular
oscillations, with each subsequent oscillatory dive during the ascent descending
less deep than the previous one by a similar depth.
The function of deep dives in R. typus and other shark species remains
unknown. One possible explanation for dives through vertical depth strata is
the acquisition of navigational cues. Although little is known about how animals navigate over long distances (Alerstam, 2006), it has been suggested that
sharks might explore the water column to gain directional information from,
for example, magnetic gradients or depth strata with different chemical properties (Klimley et al., 2002). A more likely explanation, however, is that these
dives represent search behaviour for feeding opportunities. The use of a broad
vertical habitat, extending from the surface across both epipelagic and mesopelagic zones over a time scale of hours, has also been found in plankton-feeding
C. maximus in which deep dives were suggested to relate to search patterns for
deep-water zooplankton (Sims et al., 2003). Large-amplitude changes in depth
in response to concomitant depth changes in prey concentrations have been
suggested for a wide range of marine vertebrates including all three planktivorous shark species: C. maximus (Sims et al., 2005), Megachasma pelagios Taylor,
Compagno & Struhsaker (Nelson et al., 1997) and R. typus (Gunn et al., 1999;
Graham et al., 2006; Wilson et al., 2006). The observed diel changes in preferred depth are consistent with a response to food patches that occupy deeper
depth strata during the day in vertically thermally stratified habitats (Sims
et al., 2005). In the absence of data on deep-water plankton from this region,
a related explanation may be that the recorded deep dives in the Mozambique
and Madagascar Basin represent searches through the fish’s entire vertical
niche (0–1286 m), a strategy that would enable any food layers at shallower
depths to be encountered more readily.
The archived record of shark 2’s long-distance movement across the Mozambique Channel represents the most detailed data set available on the vertical
behaviour of a R. typus in the western Indian Ocean, a key area for this species.
The long-term horizontal and vertical movements of R. typus are among the
best studied of any elasmobranch species. Some published tracks obtained from
satellite-linked transmitters have shown large geographic displacements, such as
Eckert & Stewart’s (2001) report of a R. typus swimming 13 000 km from the
Sea of Cortez to the western north Pacific Ocean and a fish tagged in the Seychelles apparently moving as far east as south of Sri Lanka (Rowat & Gore,
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J. M. BRUNNSCHWEILER ET AL.
2007). Unfortunately, these large-scale displacements are based on very few Argos position estimates, with few or no intermediate positions, and similar results could potentially be obtained from a detached tag drifting on the
surface. Tracks, including the above mentioned, lacking associated depth data
make it impossible to verify that the tag remained attached. The 1200 km longdistance movement reported for shark 2 in this study is one of the longest recorded where the geographic movement of the fish can be validated with a high
degree of certainty.
The tagged female R. typus crossed the Mozambique Channel, passed the
southern tip of Madagascar and into the Madagascar Basin as far east as
50494° E reaching the maximum value on 5 May [Fig. 1(c)]. From there,
the fish apparently turned in a north-westward direction swimming towards
the east coast of Madagascar where the tag eventually popped up on 16
May [Fig. 1(a)]. The trigger for large-scale geographic movement in R. typus remains largely unknown. A plausible explanation for planktivorous shark species is that individuals move between locations with high concentrations of
planktonic organisms (Sims & Quayle, 1998; Heyman et al., 2001; Sims
et al., 2003, 2006). Although data on plankton densities from the southern
coast of Mozambique are lacking, R. typus are regularly observed feeding in
surface waters near Tofo. The R. typus in this study was tagged in February
and initially remained in these warm and shallow coastal waters. After 20 days,
shark 2 left the coastal waters and crossed the southern part of the Mozambique Channel, an area with little primary production (Machu et al., 2005),
within 30 days before reaching the Madagascar Ridge at the beginning of April
and continuing to move eastwards into the Madagascar Basin. This area southeast of Madagascar is known to support a vast phytoplankton bloom in late
austral summer (Longhurst, 2001). Such an abundant food source might attract
planktivorous species seasonally and could explain the large-scale directed
movement of this female R. typus. This location, identified by shark 2, may represent a place where large numbers of R. typus seasonally aggregate.
We thank G. Adkison, J.-P. Botha, A. Cumming and F. Gnaegi for their efforts in
the field and J. Van Buskirk for helpful comments on an earlier version of this manuscript. J.M.B. was supported by grants from the Save Our Seas Foundation, Shark
Foundation Switzerland, Project AWARE and grant No. 11 9305/1 from the Swiss
National Science Foundation. Research by S.J.P. in Mozambique was supported by
the Save Our Seas Foundation, Casa Barry Lodge and Tofo Scuba. D.W.S. was supported by a U.K. Natural Environment Research Council-funded MBA Fellowship.
We also thank three anonymous reviewers for helpful comments on the manuscript.
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