Address for correspondence:

Bonnie J. Holmes,
Level 4,
Otto Hirschfeld Building,
The University of Queensland,
Brisbane 4072, Australia.
E-mail: b.holmes@uq.edu.au
ANTHROZOS VOLUME 25, ISSUE 4 REPRINTS AVAILABLE PHOTOCOPYING ISAZ 2012
PP. 397413 DIRECTLY FROM PERMITTED PRINTED IN THE UK
THE PUBLISHERS BY LICENSE ONLY
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Gift Giving by Wild Bottle -
nose Dolphins (Tursiops sp.)
to Humans at a Wild Dolphin
Provisioning Program,
Tangalooma, Australia
Bonnie J. Holmes*

and David T. Neil

*School of Biological Sciences, The University of Queensland,

Brisbane, Australia

Fisheries Queensland, Department of Agriculture, Fisheries and

Forestry, Brisbane, Australia

School of Geography, Planning and Environmental Management,

The University of Queensland, Brisbane, Australia
ABSTRACT Since 1992, wild dolphin provisioning has occurred on a
nightly basis at Tangalooma, a resort located on Moreton Island, Australia.
Each evening at dusk up to 12 bottlenose dolphins (Tursiops sp.) are
provided with fish in a regulated provisioning program. Since July 1998,
biologists managing the program have documented 23 occurrences of gift
giving, when several of the provisioned dolphins have offered wild-caught
cephalopod or fin fish species to staff members. The characteristics of each
of these events are presented, and we explore the relationships between
these events and their temporal patterns, and the age and sex of the dol-
phins involved. We also consider the behavioral explanations for the gift
giving, including prey sharing, play, and teaching behaviors, which have
previously been described for cetaceans and other higher mammals. Gift
giving may occur either as a discreet behavior (that may be a sequel to one
or more other behaviors such as play or food preparation), or as a part of
other behaviors, such as play and/or food sharing. It is most likely a man-
ifestation of the particular relationship between the provisioned dolphins
and the human participants in the provisioning. Gift giving has become an
established but infrequent part of the culture of the provisioned dolphins at
Tangalooma.
Keywords: bottlenose dolphin food sharing, gift giving, play, wild dolphin
provisioning
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Since 1992, Tangalooma Resort on Moreton Island, Queensland (Figure 1) has con-
ducted a wild bottlenose dolphin (Tursiops sp.) provisioning program on the beach
adjacent to the resort. In 2010, up to 11 adult, sub-adult, and juvenile dolphins would
attend the nightly provisioning sessions. In the 18 years since the programs inception, resort-
employed biologists and staff working with the dolphins have compiled extensive photographic
libraries for each animal, as well as completing detailed nightly data sheets on behavioral ob-
servations. A previously unreported behavior, observed initially in 1998, and repeated inter-
mittently up to January 2011, is that of dolphins bringing wild-caught fin fish and cephalopods
to the beach which they then offer to the resort staff members who are provisioning the dol-
phins. We refer to this behavior as gift giving, a term that will be evaluated in the conclusion
of this paper. We first provide a brief review of animal behaviors that may explain the gift giv-
ing observations. These include food sharing (e.g., reciprocal altruism, incompetent hunter)
and play behavior (including object play, predator play, and Surplus Resource Theory). We fol-
low this with a description of the setting for the gift giving events and a description of each oc-
currence. We then provide an analysis of these events in relation to their temporal patterns and
the dolphin characteristics (e.g., age, sex), and propose an explanation of these events in the
context of the behavioral theory reviewed below.
O
Figure 1. Location of Tangalooma Island Resort, Moreton Bay, Australia.
Gift Giving by Wild Bottlenose Dolphins (Tursiops sp.) to Humans
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Holmes and Neil
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Food Sharing
The origin and persistence of inter- and intra-specific cooperative behavior has been widely
discussed by evolutionary biologists. Of the published reports of intra-species cooperative
behavior, many refer to food sharing. Intra-specific food sharing behavior is well documented
and common in non-human animals (for examples, see Vahed 1998, for insects; Tryjanowski
and Hromada 2005, birds; Hauser et al. 2003, small mammals; Connor and Norris 1982,
cetaceans; and Rose 1997, primates). However, food sharing that occurs between non-related
adults that are neither closely related nor reproducing together apparently challenges
evolutionary theory, because explanations like inclusive fitness or courtship feeding are not
applicable (de Kort, Emery and Clayton 2003). One popular explanation for these behaviors is
that food sharing is maintained through reciprocal altruism (reciprocity) (Trivers 1971), where
food is shared because the receivers are likely to reciprocate at a later date (Scheid, Schmidt
and Noe 2008), which is ultimately selfish. This has been described for chimpanzees (de Waal
1997), capuchin monkeys (Hauser et al. 2003), and vampire bats (Wilkinson 1984).
Several studies describe cetacean behaviors at sea and in captivity that have been re-
peated often enough that some assessment of them can be made (Connor and Norris
1982). One trait, often mentioned, is that dolphins and other toothed whales show altruistic
behavior toward both conspecifics (intra-species) and cetaceans of other species (inter-
species). Johnson (1982) reported food sharing behavior in captive Amazon river dolphins
(Inia geoffrensis), where an adult male was observed routinely sharing pieces of fish with
tank mates. When the health of one of the adult females worsened (due to a pneumonic
condition), he offered her whole fish. In 1999, wild rough-toothed dolphins (Steno
bredanensis) in Costa Rica were first observed sharing large mahi mahi (Coryphaena
hippurus). Three dolphins took turns in carrying and chewing on the fish before it appeared
to be returned to the original keeper of the fish (Pitman and Stinchcomb 2002).
Fedorowicz, Beard and Connor (2003), also in Costa Rica, reported two bottlenose dol-
phins sharing a fish, where possession of the fish changed 11 times during a 30-minute ob-
servation period. They conclude that the repeated changes in possession of the food item
was, at best, extremely rare in other taxa.
Inter-species food sharing is more rarely observed and more difficult to explain, given that
reciprocation would not be expected to occur and inclusive fitness would not be served. Mor-
ris (1986) describes a commonly observed behavior of food sharing between domestic cats
and their human owners. The cats that most commonly bring home prey as gifts for their own-
ers are spayed females. Morris suggested that their instinct to bring food home for their kit-
tens is redirected to their human companions, who they view as incompetent hunters. Under
natural conditions, a mother cat introduces her kittens to prey very gradually by first bringing
home dead prey and eating in front of them. Next she brings home live prey and shows them
how to subdue and kill it. After many lessons, the kittens will have acquired the necessary
skills to leave the nest and obtain their own meals.
Similar observations from wild populations are rare. One such interaction, between a wild
dolphin and a human, was described by Connor and Peterson (1994). National Geographic
photographer Flip Nicklin was in Hawaii photographing false killer whales. They reported that
one swam up to him with a large mahi mahi in its mouth. The whale released the fish in the
photographers face and backed away. Nicklin accepted the fish, then returned the fish to the
whale, who accepted it before swimming away. Whether this was play behavior by the whale,
or whether altruism by the whale led it to share its food with the photographer, is unclear.
!" $%&' ()*+,'-./0&12345 6 678+86( (06? :; :1<= >??
In 2009, a similar occurrence was reported by National Geographic photographer and bi-
ologist Paul Nicklen, who was photographing leopard seals in Antarctica. On his first day in the
water he encountered an adult female leopard seal hunting penguins on a nearby ice shelf. The
seal was quick to investigate the camera and photographer by mouthing them before catch-
ing a whole live penguin and pushing it to the camera, which Nicklen suggested the leopard seal
saw as his mouth. The seal then released the penguin which swam away. Nicklen described
the leopard seals demeanor as one of confusion that this useless predator in her ocean was
probably going to starve to death. Nicklen reported that she then became quite panicked and
proceeded to bring him weak penguins, then dead and partially consumed penguins, with the
seal making a show of how to eat them in front of the photographer. This behavior continued
each day for the remaining four days of filming with the seal (Richard 2009).
1
Play Behavior
Animal play behavior is well documented (Fagen 1981; Smith 1982; Martin and Caro 1985;
Barber 1991; Caro 1995), and may be defined by its attributes, by operational means, by ex-
ample, by enumeration of behaviors called play, or by contrast with similar behaviors (Fagen
1981). Many descriptions of play in mammalian species are anecdotal, with few researchers
able to include quantitative data suitable for testing theory. However, even anecdotal accounts
serve to illustrate the range, variety, and coherence of animal play and bring long-neglected,
theoretically significant phenomena to light (Fagen 1981), particularly for wild animals to which
relatively little empirical research has been devoted (Bekoff 2000).
Mammals, including primates, typically play 110% of the time (Fagen 1981). Percentages
vary significantly between adult and juvenile animals, with juveniles playing more frequently
and for longer periods. The main hypotheses are that juvenile play may serve as practice for
predatory behavior (in predator species), or for tool use (particularly in primates); and may en-
able exploration of stimuli, objects, and environments new to the juvenile (Hall 1998; Gibson
and Mann 2008). Adult play is less commonly observed and is perhaps more resistant to ex-
planation. It is also likely that the motivations and function of adult play behavior are distinctly
different from its juvenile counterpart (Hall 1998).
Animal play behavior is predicted to occur more commonly in those animals which are not
constantly near their physiological limits, which are not in nutritional stress and which have
large body size (reducing the cost per kilogram of locomotion), encapsulated by Surplus
Resource Theory (Burghardt 1988). Burghardt also suggested that well-cared for captive
animals could be expected to play more than wild animals since they are more likely to have
the surplus energy required to spend time playing. Extended parental care also gives juveniles
time for boredom play (Hall 1998).
Involvement of inanimate objects in various kinds of animal play activities is known as
object play. In adult animals it is usually difficult to associate with any particular function.
Some studies with domestic animal subjects have provided evidence for the relationship be-
tween object play and predatory behavior including practice predatory play with the prey and
normal predation behavior, for example, the kill bite (Hall 1998). Rasa (1971) found that North-
ern elephant seals preferred play with small, moving objects and suggested that this might be
explained by the animals search image for small moving prey (fish and squid). In predatory
behavior, any objects (including prey) which possess particular stimulus characteristics asso-
ciated with prey, such as small size and complex and rapid movement, attract the predator and
can elicit play as well as predation (Hall 1998).
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Differentiating play behavior from other behaviors, particularly when prey items are involved,
is difficult. For example, in 1999 wild rough-toothed dolphins in Costa Rica were observed
sharing a large mahi mahi (Coryphaena hippurus). Three dolphins took turns in carrying and
chewing on the fish before it appeared to be returned to the original keeper of the fish, mak-
ing it difficult to determine if the dolphins were planning on eating the fish, or if they were just
playing with it (Pitman and Stinchcomb 2002).
Play behavior between individuals of different species has been documented in wild pop-
ulations (see Fagen 1981) and, more frequently, for animals in captivity. For interspecific play,
individuals must recognize and respond to another species play signals. Many animals in
captivity are kept for educational and entertainment purposes, and undergo training through
positive reinforcement, usually with food items. It is these food items, or other items provided
in the aquaria (i.e., toys) that have been used by captive bottlenose dolphins to initiate play
with their trainers, which often extends the interactive session with their human trainers (Dima
et al. 2008).
The terms food sharing, reciprocity, and cooperation are well defined in the literature.
As the events at Tangalooma do not appear to be entirely consistent with any previously
described behaviors, we refer to the phenomenon described below as gift giving. Following
is an account of the Tangalooma dolphin provisioning program and of the gift giving events
which adult, sub-adult and juvenile wild bottlenose dolphins (Tursiops sp.) have initiated.
The Tangalooma Wild Dolphin Provisioning Program
The establishment of the wild dolphin provisioning program at Tangalooma is well documented
(e.g., Green and Corkeron 1991; Orams 1994; 1995; 1996; Neil and Brieze 1998). The provi-
sioned dolphins consist predominantly of two matrilines (Figure 2) and, at January 2011, in-
cluded six adults (four male and two female), two sub adults (one male and one female), and
three calves. The sub-adults are recently weaned calves that were born in August and
September, 2004. In late 2008, Tinkerbell, one of the provisioned dolphins, gave birth to
another calf, Phoenix (female), and Shadow, another mature provisioned female (now
presumed deceased), also gave birth to her second calf Zephyr (male). In December 2010,
Tangles gave birth to her first calf (unnamed) whose sex is unknown.
Resort staff influence wild dolphin behavior by both positive and negative reinforcement
with high quality teleosts, snap frozen after capture. No physical contact with the dolphins is
permitted by staff or guests. Resort staff members begin each nightly session with a staff-
only feed to ensure the dolphins are positioned in the correct area for provisioning. No guests
accompany the staff into the water at this time. Following the staff feed, two guests accom-
pany each staff member into thigh-deep water to hand feed two dolphins in rotation until all
guests have participated. Staff and guests all enter and depart the water at the same time.
In addition to those guests who participate in the provisioning, an additional 250 guests
(approx.) are able to observe the provisioning and hear an educational commentary from a
seating area on the resort jetty immediately adjacent to the provisioning area. The provision-
ing is organized into several lanes, each with one staff member, two guests, and two dol-
phins. At the time of the last recorded gift giving event, Echo (18-year-old male) and Nari
(13-year-old male) were fed in lane 1; Tinkerbell (20-year-old female) was fed in lane 2. Lane
3, until recently, fed Shadow (16-year-old female) and Tangles (10-year-old female); with Rani
(~18-year-old male) and Bobo (~24-year-old male) fed in lane 4. The two youngest sub adults,
Storm (male) and Silhouette (female), are 6 years of age and were provisioned in lane 5.
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Figure 2. Age, sex, and matrilines of dolphins attending the Tangalooma provisioning
program. Shading indicates deceased dolphins.
Calves born to provisioned mothers are not fed until 3 years of age. Provisioning is
undertaken under a permit issued by the Queensland Department of Environment and
Resource Management.
Methods
Although the program has been in operation since 1992, it is only over the past 13 years (July
1998January 2011) that 23 cases of prey items being caught by the dolphins and then of-
fered to resort staff at the Tangalooma provisioning program have been documented. For the
purpose of this paper we have called this phenomenon gift giving. Accounts of these
events are included in reports completed each night during and immediately after the pro-
visioning by resort biologists and other trained staff members, and photographs taken and
archived when possible. Historical records of the gift giving observations are largely anec-
dotal. The majority of the staff that record information on the nightly data sheets are not be-
havioral scientists. Each record of gift giving behavior was assessed by the authors for
reliability by contacting current and past staff members for independent corroboration of
events. This was possible for 20 of the 23 cases. The authors were present for the most re-
cent event in January 2011. Details of each of these events are outlined in chronological
order below (summarized in Table 1).
Gift Giving by Wild Bottlenose Dolphins (Tursiops sp.) to Humans
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Results: Observations of Gift Giving Events
Details of each of these events are outlined in chronological order below and summarized in
Table 1.
July 2, 1998Fred
At 19:05, when the guest feed had finished, Fred captured a moray eel (Gymnothorax sp.)
which was seen attempting to bite at Freds head. Fred killed the eel and offered it to staff
member #1 at the final staff-only feed (Table 1). The eel was returned to Fred, who then
departed in a south-westerly direction with the eel, the fate of which is unknown.
August 25, 1998Tinkerbell
Tinkerbell arrived at 18:15 with a 50 cm long flathead (Platycephalus fuscus) in her mouth.
Throughout the program, rather than stationing and receiving fish handouts from guests, she
swam around the offshore end of the feed area attempting to break off the fishs head, with
eventual success. Unusually, Tinkerbell (provisioned in lane 2) swam to lane 3 and offered the
flathead to staff member #2. Tinkerbell then refused to accept any fish, including the flathead
that was offered back to her by staff in lane 2. The flathead was retained, and is now preserved
and on display at the resort.
August 29, 1998Echo
At the commencement of the feed, Echo brought in a live porcupine fish (Diodon sp.)
approximately 20 cm long. He offered the fish to staff member #3 who held up the fish and
showed it to the guests watching from the jetty. The fish was then returned to Echo, who
swam west out of the feed area with the fish, the fate of which is unknown.
April 13, 1999Shadow, Nari, and Rani
Prior to commencement of provisioning, Rani was seen playing with a live eel. During the feed,
Nari and Shadow each caught a rabbitfish (Siganus sp.) and offered them to the staff feeders
in their respective feeding lanes. The fish were returned to the dolphins and the fate of the fish
was not observed.
Rani and Echo caught and killed an eel (Gymnothorax sp.) that Rani was seen playing with
earlier. Rani then took the eel to staff member #3, who held up the eel to show guests and have
photographs taken. The eel was returned to Rani who departed the provisioning area with the
eel. Its fate is unknown.
November 21, 2000Shadow
At 19:15, Shadow caught a large striped butterfish (Selenotoca multifasciata). She was
observed at the offshore end of the feed area trying to break up the fish on the sea floor.
Shadow broke off the fishs head and then approached the beach with the fish. She offered
the fish to staff member #6. Shadow would not take the butterfish back when it was offered
to her. The butterfish was retained and is now preserved and on display at the resort.
June 11, 2001Shadow
During the provisioning Shadow was seen to catch a rabbitfish (Siganus sp.) which she killed
shortly after. The fish was then offered to staff member #7. The fate of the fish was not recorded.
July 9, 2001Rani
During the provisioning program Rani was seen to catch a rabbitfish (Siganus sp.). The fish was
then offered to staff member #8. Fish kept and preserved.
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October 7, 2001Nari
During the provisioning, Nari caught a rabbitfish (Siganus sp.) and brought it to staff member
#3. The staff member held the fish when Nari offered it to her, but he refused to let go, seem-
ingly playing a tug of war. After the staff member released the fish, Nari swam out of the feed
area in a north-westerly direction. The fate of the fish was not observed. He returned five min-
utes later and was provisioned as normal.
October 10, 2001Rani
At 19:10 Rani brought in a rabbitfish (Siganus sp.), and offered it to staff member #9. The staff
member did not accept the fish (knowing they have poisonous spines) and Rani played with
the fish for the next five minutes before consuming it.
April 4, 2002Shadow
Prior to commencement of provisioning, Shadow was observed hunting around the jetty pilings.
She entered the feed area with an octopus (Octopus sp.) in her mouth, which she appeared to
show to Tinkerbell and Tangles before shaking the octopus vigorously. At the commencement
of provisioning, Shadow dropped the octopus to receive a fish thrown from the jetty by staff,
then retrieved it from the seabed. When staff entered the water Shadow offered the octopus to
staff member #8. Shadow accepted a fish, and the octopus was then returned to Shadow, who
again shook it before taking it into deeper water and dropping it. Each time staff and guests were
in the water Shadow would return to the shallows to feed and then return to the octopus in
deeper water. Shadow had consumed the octopus before the end of the feed.
April 26, 2003Rani
During the provisioning program Rani caught a rabbitfish (Siganus sp.), killed it, and offered it
to staff member #9. The fish was then returned to Rani who ate the fish. He fed as normal for
the remainder of the program.
January 18, 2006Silhouette
Silhouette arrived at the program with her mother Shadow as usual. During the program
Silhouette caught a pufferfish (Tetractenos sp.) and offered it to staff member #10 in lane 3,
where Shadow was provisioned. Silhouette held the pufferfish in her mouth and nudged the
staff members hand. The staff member then took the fish and let it swim away. Silhouette
then swam and caught the pufferfish again, and returned to the staff member, offering it again.
This occurred five times until Silhouette appeared to lose interest, and the pufferfish swam
away. At the time Silhouette was 17 months old and not yet provisioned.
May 27, 2006Shadow and Fred
Shadow arrived, swam around the feed area, and departed before returning shortly after with
a large long finned eel (Anguilla reinhardtii). When the program commenced, Shadow stationed
in her normal lane 3 position with the eel and did not accept any fish until staff member #11
took the eel, which Shadow freely released. Shadow was then given several of the provision-
ing fish before the eel was returned to her. The staff and guests then left the water while
Shadow swam around with the eel. When the staff member returned to the water with two
more guests, Shadow again brought the eel in and relinquished it in order to accept fish. This
continued for several provisioning rotations until Shadow would not accept the eel when it was
returned to her and it floated out into the feed area. Fred then picked up the eel and offered it
to staff member #12 in his regular provisioning lane 2. The staff member took the eel from Fred
after he released it into her hand. The eel was retained and preserved.
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May 28, 2006Shadow
Shadow entered the feed area at 17:51 with a long finned eel (Anguilla reinhardtii). A similar
sequence of events occurred as on the previous night. However, after several rotations, when
staff member #11 again returned the eel, Shadow tail slapped the water and refused to accept
it. The eel was retained and preserved.
December 1, 2006Tinkerbell
At 19:01 Tinkerbell was observed with a half eaten tuna (Thunnus sp.) in her mouth and, over
the following five minutes, eating portions of the fish that she had torn off. She then repeat-
edly dropped the fish and circled it before picking it up again. At the commencement of the
feed (19:21), Tinkerbell dropped the fish at the feet of staff member #14 and left it there for the
duration of the feed. After the final feed the tuna was offered back to Tinkerbell, who then
backed away from it. She then took a few more of the provisioned fish before the tuna was
offered again, but she refused to accept it.
June 4, 2007Silhouette
At 18:50, during the last (staff only) feed for the evening, Silhouette (now 3 years old and
provisioned) left her station and swam quickly toward the back of the feed area. She returned
quickly to the provisioning station with a squid in her mouth and was seen eating it. She then
dropped half of the squid in front of the staff member #2. The feeder gave it back to Silhou-
ette who again dropped it at the feeders feet. The squid was again returned to the dolphin,
who used her rostrum to push the squid back into the feeders hand. This behavior occurred
several times until the squid was finally returned to the dolphin and the staff member left the
water. No guests were in the water at the time.
June 18, 2007Shadow
At the commencement of the feed (18:15), Shadow arrived with a half eaten tuna (Thunnus
sp.) and was observed shaking her head with the tuna in her mouth. During the first staff-only
feed, Shadow dropped the partially eaten tuna in the hand of staff member #15. The feeder
tried to return it to Shadow by placing the tuna under the water, but both Shadow and Tan-
gles (who feeds in the same lane) backed away. The staff member then took the tuna out of
the water and both dolphins moved back into their provisioning positions. Photographs were
taken and the fish carcass discarded.
June 23, 2008Rani
Before the provisioning program commenced, Echo was observed chasing an eel in the shal-
lows on the south side of the jetty, adjacent to the feed area. During the first staff feed, Rani
had the now dead eel in his mouth and appeared to be playing with it. On the second feed,
staff member #14 (unaccompanied by guests) entered the water and Rani stationed to accept
fish. He dropped the eel at the feeders feet. When the eel was offered back to Rani, he did
not accept it. The eel floated near his head until he nudged it back to the feeder again. This
occurred several times, with Rani flipping the eel out of the water towards the staff feeder on
one occasion. At the end of the evening during the last staff feed, Bobo (who stationed with
Rani in lane 4) picked up the eel and was last seen swimming away with it, with Rani follow-
ing. At the final feed, when fish are thrown from the jetty, Rani returned to accept a fish, but
Bobo did not. The fate of the eel is unknown.
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July 12, 2008Tinkerbell
During the feed, Tinkerbell and Shadow left the feed area for approximately five minutes. Tin-
kerbell then returned swiftly, stationing in her usual position with a large (4 kg) tuna in her mouth,
held by the tail. She released the fish in front of staff member #16, who then picked it up. The
tuna had had the head and some of the body removed. After several provisioning fish were
given to Tinkerbell, the tuna was offered back to her. Tinkerbell would not take the tuna and
moved to station in another lane. When the tuna was removed from the water, she returned
to her normal provisioning position. At the cessation of the feed, the staff offered the tuna back
to her again four times. Tinkerbell did not accept the tuna, and was observed shaking her
head vigorously from side to side.
January 5, 2011Tangles
Tangles arrived at the provisioning area at 19:04 with her 3-week-old calf. At 19:08 Tangles
caught a large butterfish and was observed repeatedly releasing it and then catching it again.
At 19:13 Tangles moved to the shallows holding her fish whilst her calf was nearby. The calf then
swam behind her into deeper water and Phoenix (Tinkerbells older calf) was observed swim-
ming to the calf and accompanying it into shallower water again. At 19:22 Tangles was ob-
served tearing at the butterfishs head, whilst her calf swam behind her with Phoenix. During the
staff-only feed, Tangles relinquished the butterfish to staff member #17 to receive a provisioned
fish. The staff member then offered the fish back to Tangles, who would not accept it back. The
butterfish was kept, photographed, weighed, and preserved by resort biologists.
Discussion: Characteristics of the Dolphin Gift Giving
Gift Recipients
All except one case of food sharing involved the dolphins giving the food item to the staff mem-
ber in the lane in which the dolphin was normally provisioned. This behavior is expected, given
that the dolphins are, through positive and negative reinforcement, only provisioned fish in a
specific lane to minimize aggression and encourage routine and cooperation through learned
behavior. Each dolphin learns that they will only be provisioned fish in their allotted lane and, if
food sharing was performed in order to exchange fish, the dolphins would know that this
would be unsuccessful in another lane.
The exception to the aforementioned pattern occurred in August 1998 when a large dusky
flathead was gifted by Tinkerbell. Tinkerbell (8 years old at the time) was usually fed in lane 2,
but swam to feeder #2 who was working in lane 3. She dropped the flathead in front of feeder
#2 and swam back to her usual lane to receive a fish. Feeder #2 was a long standing staff
member of the resort, who was involved in the development of the program from its inception.
Dolphins, in both captive and semi-captive settings, are known to have the ability to distinguish
between individual trainers (Thieltges et al. 2011) and also exhibit preferences for trainers
based on their behaviors and compliance when working together (Norris 1974). Although reg-
ular staff changes occur and the dolphins receive fish only in pre-determined lanes, at least in
this case the dolphin exhibited a preference for a specific staff member.
Temporal Patterns
Gift giving frequency varies both secularly and seasonally. There was a lag of six years be-
tween commencement of the provisioning program (1992) and the first occurrence of an item
being offered by a dolphin (by Fred in July 1998). There are anecdotal reports of the Tanga-
looma dolphins engaging in play behavior with inanimate objects (sticks, seaweed, plastic
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bags) during the programs developmental phase (M. Orams, personal communication), al-
though there is no record of the dolphins offering or giving these objects to resort staff at the
time. The time required for the dolphins to establish both trust of, and some form of relation-
ship with, the humans involved may provide a partial explanation for this lag period. The long
duration of this lag time (compared with, for example, bottlenose dolphins introduced into cap-
tivity) may be a consequence of the short period of interaction between the dolphins and hu-
mans (c. 1hr/day) and the prohibition of any physical contact (e.g., touching, stroking) between
the humans and the dolphins. For the 13 years subsequent to this initial six-year lag, gift giving
has occurred at the rate of approximately 1.8 events per year, albeit with significant gaps in
occurrence (May, 2003December, 2005; August, 2008December, 2010).
Lactating females consume between 50% and 100% more food than quiescent (neither
pregnant nor lactating) females and mature males (Cockroft and Ross 1990). Neither Tinker-
bell nor Shadow (collectively responsible for 35% of all gifts) engaged in any gift giving whilst they
were lactating with their young. This provides a partial explanation for the lack of gift giving
events from 20002003 for Tinkerbell (with calf Tangles) or from 20042006 for Tinkerbell (with
calf Storm) and Shadow (with calf Silhouette). Similarly, Bess was pregnant and lactating with
Rani ~19911994 and with Nari ~19961999, and Beauty with Shadow in ~19931995
(Beauty died 1995). This is consistent with their requirement for higher food intake to supple-
ment milk production for their young and, at these times, they probably do not readily part with
any prey items caught, regardless of nutritional value. It is also consistent with Surplus Resource
Theory, that is, that animals not near their physiological limits will exhibit more play behavior
than those who are. Thus, temporal patterns of gift giving, including the lag period before gift
giving commenced, seem strongly associated with the reproductive status of the adult females
in a manner consistent with Surplus Resource Theory.
There is strong seasonality in the occurrence of the gift giving events, with the majority oc-
curring in winter (12) and autumn (5), with relatively few in spring (3) and summer (3). Seasonal
changes (e.g., in water temperature and fish spawning) affect the species richness and abun-
dance of prey items in Moreton Bay, with peak abundance recorded in the summer months (see
Young and Wadley 1979; Blaber and Blaber 1980; Burgess 1980; Quinn 1980; Morton, Pollock
and Beumer 1988). Relatively high prey abundance in summer is a possible explanation of the
lower attendance of dolphins at the provisioning program during those months. Lower gift giving
rates in summer are expected, given the lower attendance rates. However, summer attendance
is 78% of winter attendance, whereas summer gift giving is only 25% of winter gift giving,
suggesting that the strong seasonality of gift giving is not explained by seasonality of dolphin
attendance alone and is inversely related to the seasonal abundance of prey.
Seasonal mating behavior may also affect temporal patterns of dolphin gift giving. Six
calves have been born to mothers provisioned at Tangalooma, and all of these births occurred
between August and January. Given that bottlenose dolphins gestate for approximately 12
months, it follows that mating activity is greatest during these months for this group of dolphins.
During summer, the breeding adult females that attend the program (Tinkerbell, Shadow and
more recently Tangles) are often joined by groups of males that will swim and station behind
the females whilst they are provisioned, often also engaging in harassment and sexual ac-
tivity. This usually occurs for the duration of the feed and continues when the females depart
the provisioning area each evening after provisioning ceases. This herding and harassment
behavior by males is well documented elsewhere (see Connor, Smolker and Richards 1992;
Connor, Heithaus and Barre 2001; Mller et al. 2001). The high level of social and sexual
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activity during spring and summer may influence the frequency of gift giving, where energy
consumed during courtship and mating behavior may reduce surplus resources, and leisurely
play behavior is at a minimum. Thus, it seems likely that the strong seasonal pattern of gift giv-
ing is influenced by several factors including dolphin attendance at the provisioning program,
mating behavior, and inversely to prey availability.
Sex, Genetic Relationships and Age
Shadow (female) was responsible for giving 30% of the prey items offered (seven in total). Rani
(male) was responsible for 22% (five), and Tinkerbell (female) 13% (three). Silhouette, Fred, and
Nari have given two prey items (9%) each since 1998. Echo (male) and more recently Tangles
(female) have both given one prey item (4%). About half of the gift giving events were by female
dolphins (57%) and half by males (43%). The first six gift giving events were carried out by six
different dolphins, four males and two females, over a ten-month period.
Most of the dolphins provisioned at Tangalooma are of two matrilines (Figure 2), derived from
Beauty (died 1995) and Bess (died 2000). Although neither Beauty nor Bess participated in gift
giving (perhaps due to their reproductive status), both of Bess offspring (males) have offered fin
fish items, as have Beautys offspring (Tinkerbell and Shadow, females). Shadows offspring
(Silhouette; female; 3 years old) has shared two food items, and only recently has one of Tinker-
bells two offspring (Tangles; female; 10 years old) participated in the gift giving behavior.
With the exception of Fred, whose first offering was the first gift giving event recorded, and
Tangles recent event at 10 years of age, all of the dolphins who have given a gift were aged
8 years or younger at the time of their first offering, and the gift giving has generally contin-
ued into adulthood (Table 1). Exceptions are Nari (gifts at age 2 and 4 years), now 13 years
old, and Echo (at 6 years), now 18 years old, who have not subsequently participated.
Silhouettes precocious gift giving at the age of one year is noteworthy. This is not unexpected,
given that juvenile play may serve as practice for predatory behavior (Hall 1998). Play among
the adult dolphins, which occurred in association with four of the 23 events, may be explained
by some combination of play per se, play while teaching juveniles how to forage and prepare
prey items to consume, and/or play enabled by surplus resources available due to food
supplied by the provisioning.
Fish Species and Quality
The species of prey items gifted since 1998 range from low value food items (such as the poi-
sonous porcupine fish and rabbitfish) to high value oily fish and cephalopods (such as tuna
and octopus) (Table 1). In general, animals should prefer prey types which provide the great-
est profitability, that is net energy return per unit time spent handling (Stephens and Krebs
1986). Dolphins have the capacity to exhibit rational choice of food items, probably based on
relative energy obtained from the prey item and individual dolphins exhibit considerable varia-
tion in prey item choice (Barros and Odell 1990; Dill, Dill and Charles 2003). It is clearly more
profitable for a dolphin to give up a low value prey item for a high value one, perhaps even if
the energy spent to obtain the item was significant. The willingness to give both low- and high-
value prey items may be explained by the dolphins strong preference for the silver biddies
(Gerrus ovatus) that are provisioned in the program. At the inception of the Tangalooma pro-
visioning program in 1995, prey item preferences were tested with a range of local species in-
cluding mullet (Mugil georgi), silver biddies, winter whiting (Sillago maculate), mackerel
(Scomber australasicus), grinners (Saurida sp.) and yellowtail (Trachurus novaezealandidae)
(Orams 1995), with silver biddies being the most popular choice made by the dolphins.
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The value of the fish items offered and the manner in which they were offered varies be-
tween events. All of the high value teleost items (three tuna, two butterfish, and one flat-
head), were large items which had their heads removed by the dolphins prior to being gifted
to the staff members. Many dolphins and sea lions remove heads from large fish before con-
suming them, and bottlenose dolphins will strip flesh from spiny fish (Perrin, Wrsig and
Thewissen 2008). Odontocetes cannot chew prey and must spend time handling large prey
items. For example, bottlenose dolphins will drag large fish along sandy bottoms until pieces
are small enough to swallow or are broken off (Perrin, Wrsig and Thewissen 2008). In the gift
giving cases, it may be speculated that the dolphins were preparing the prey item as food,
whether to eat themselves or to give to a human. The dolphins may also be engaging in a com-
bination of play and/or predatory play, demonstrating to young how to catch and prepare large
meals. The observation of Shadow showing off her octopus catch to other dolphins in the
group may also represent a status enhancing display. Thus, although there are several pre-
cursor behaviors (and possible behavioral motivations) that occur separately or in combination,
at some point the dolphin appears to make a choice to relinquish the high value food item and
give it to a human, with the opportunity to do this perhaps underpinned by the resource surplus
provided by the provisioning.
The other fish species involved in the gift giving events are what could be regarded as low
quality food items including seven rabbitfish, six eels, and two toad/porcupine fish (both of
which were still alive). The low value prey items were not prepared for consumption as de-
scribed above, but were played with before being offered to the staff members. In some
cases, possession of the fin fish also changed between dolphins during the feed, consistent
with the observations of Pitman and Stinchcomb (2002) for wild rough-toothed dolphins.
The first gift given by Silhouette warrants special mention. This case, where the live
pufferfish was caught and returned to the feeder several times, is consistent with play be-
havior. It is not consistent with food sharing, reciprocity, or the incompetent hunter ideas.
The pufferfish game, where the small fish are caught and, when expanded due to stress,
repeatedly flung into the air and retrieved by the calves has frequently been observed at
Tangalooma (T. Hassard, personal communication). This form of play is vital in Silhouettes
foraging development as she grows to adulthood (Sargeant and Mann 2009). At Tanga-
looma, calves are not normally provisioned until they reach three years of age, in order that
they develop natural hunting skills and avoid any risk of dependence on provisioning. At age
1 year, Silhouette had never been provisioned or included in the program in any way, nor
had she witnessed another gift giving event by one of her provisioned conspecifics, which
precludes the idea of direct social learning (Laland and Hoppitt 2003). A more appropri-
ate explanation may be that there is a form of learned trust of the staff members at
Tangalooma, so that Silhouette, through observation of the provisioning of her mother (and
other members of the group), was confident in initiating play with the staff member, notably
in lane 3 where her mother was provisioned. Learning from observation of, or interaction
with others, allows animals to match the demands of their environments quickly (Galef and
Giraldeau 2001; Laland 2004), and is considered to be a critical underpinning of complex
cognition, behavioral flexibility, and individual differences (Whiten and van Schaik 2007;
Sargeant and Mann 2009). Social learning is also considered the most basic requisite for
culture in animals (Laland and Hoppitt 2003). Thus, although social learning cannot
explain Silhouettes precocious gift giving, it may explain the behavioral context which
facilitated it.
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Gift Giving by Wild Bottlenose Dolphins (Tursiops sp.) to Humans
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The dolphins at Tangalooma have been provisioned with the same species of fish each
evening for many years, including the six years prior to the first gift given by a dolphin. This sug-
gests that the dolphins do not view the gifted item as an exchange of food, and therefore the
idea of reciprocity seems not applicable because the dolphins know that they will be provi-
sioned regardless of whether or not they provide a gift. For this reason, it is also unlikely that
the dolphins view their human counterparts as incompetent hunters as described by Morris
(1986) in cats. The account given by Paul Nicklin and the leopard seal in Antarctica more ap-
propriately fits the incompetent hunter idea than the gift giving that has occurred at Tanga-
looma, with Nicklin describing the leopard seal viewing him as ... this useless predator in her
ocean Perhaps had he reciprocated in kind with a fish item (or penguin), her behavioral
response to him would have been different.
The behavioral mechanisms behind the dolphin gift giving phenomenon are difficult to as-
certain. We believe that, given the diversity of dolphin ages and sex and the various prey types
given, there is a range of behavioral motivations that drive the gift giving phenomenon at Tan-
galooma. Low value prey items are generally played with prior to the gift giving events. High
value prey items appear to have stronger elements of predatory play, which may be a function
of teaching the younger dolphins how to hunt and subdue the prey, and prepare it for con-
sumption by removing the fishs head. We argue that these high value items were initially seen
as prey because they were prepared for consumption, but were then given to staff members
instead, and often not accepted back. We also recount Shadows showing off behavior
which, while it appears playful, may be important in the establishment of social standing within
the group. The abovementioned behaviors may occur separately, or in combination, as a
precursor to the actual gift giving events, but are not part of the act of gift giving.
Given this pattern of behaviors and events, it appears that the gift giving: (a) is not associ-
ated with the dolphins attempting to enhance nutritional intake by reciprocity; (b) is somehow
associated with the various manifestations of play behavior by the dolphins; and (c) is a dis-
crete behavioral type which is probably largely explained by some undefined aspect of the re-
lationship between the provisioned dolphins and the human participants in the provisioning.
Although the gift giving is infrequent (~1.8 events/year) (and records of it are anecdotal reports
by, in most cases, non-behavioralists) it has been carried out seven times by one dolphin
(Shadow), and by eight different dolphins in total.
A gift is, by definition, an item which is given, in this case by the dolphins to the hu-
mans. Occurrences may or may not be associated with particular behavioral precursors or
with intentions of food sharing or expectations of reciprocity. Behavioral explanations differ
among the gift giving events recorded but, regardless of the explanation, the term gift giv-
ing would appear valid in each case. It is most likely a manifestation of the particular rela-
tionship between the provisioned dolphins and the human participants in the provisioning.
Gift giving has become an established but infrequent part of the culture of the provisioned
dolphins at Tangalooma.
Acknowledgements
We thank the staff at Tangalooma Wild Dolphin Resort for their continued support and providing
us with the necessary data and time with the dolphins in order to complete this paper. We
would also like to thank Dr Celine Frre for comments on an earlier draft, and Dr Mark Orams
and one anonymous reviewer for their time and effort in providing detailed and helpful
comments on the final manuscript.
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Note
1. Video downloaded and available from corresponding author upon request.
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