Marine Biology (2004) 145: 435–444
DOI 10.1007/s00227-004-1339-1
R ES E AR C H A RT I C L E
K. J. Benoit-Bird
Prey caloric value and predator energy needs: foraging predictions
for wild spinner dolphins
Received: 17 April 2003 / Accepted: 16 February 2004 / Published online: 25 March 2004
Ó Springer-Verlag 2004
Abstract Spinner dolphins (Stenella longirostris) feed on
individual small (2–10 cm long) prey that undergo diel
vertical migrations, presumably making them inaccessi-
ble to dolphins during the day. To examine how time,
prey behavior, prey distribution, and energy needs
constrain dolphin foraging, a calorimeter was used to
measure the caloric content of prey items. These data
were combined with information on prey distribution in
the field and the energetic needs of dolphins to construct
basic bioenergetic models predicting the total prey con-
sumption and mean feeding rates of wild dolphins as
well as potential prey preferences. The mean caloric
density of mesopelagic animals from Hawaii was high
(2,837 cal/g wet weight for shrimps, squids, and myc-
tophid fishes). Their total caloric content, however, was
low because of their small size. Energy value of prey and
energetic needs of spinner dolphins were used to exam-
ine the effect of time and energy constraints on dolphin
foraging. The results predict that spinner dolphins need
to consume an estimated minimum of 1.25 large prey
items per minute to meet their maintenance energy
needs. If the additional energy costs of foraging are
considered, the estimated necessary foraging rate is
predicted to increase only slightly when large prey are
consumed. If smaller prey are consumed, the total en-
ergy demand may be twice the basic maintenance value.
Prey density and size are predicted to be important in
determining if dolphins can forage successfully, meeting
their energetic needs. The prey size predictions compare
well with results from previous gut content studies and
from stomach contents of a recently stranded spinner
dolphin that had enough prey in its stomach to meet its
estimated basic maintenance energy needs for a day.
Communicated by P.W. Sammarco, Chauvin
K. J. Benoit-Bird
Hawaii Institute of Marine Biology,
P.O. Box 1106, Kailua, HI 96734, USA
E-mail: benoit@hawaii.edu
Tel.: +1-808-2475063
Fax: +1-808-2475831
Finally, the results suggest that spinner dolphins are
time and therefore efficiency limited rather than being
limited by the total amount of available prey. This may
explain the diel migration exhibited by spinner dolphins
that allows them to follow the movements of their prey
and presumably maximizes their foraging time.
Introduction
Bioenergetics models of foraging have provided useful
predictions about the behavior and population dynamics
of many predators. For example, models incorporating
prey energy values and predator energetic needs have
been used to predict dive times in blue whales (Acevedo-
Gutierrez et al. 2002), total hunting time for wild dogs
(Gorman et al. 1998), and growth rates in flounder
(Burke and Rice 2002). Models incorporating foraging
energetics in pelagic predators have also been useful in
examining prey preferences and understanding the
impact of the predator on the ecosystem (Bunce 2001).
These kinds of predictions are particularly useful when
studying populations that are difficult to observe
directly, such as pelagic marine animals. These models
provide testable hypotheses for data that are often more
easily measured than the parameters of the model and
can also provide explanations for observed behaviors.
Spinner dolphins, Stenella longirostris, a subtropical
pelagic species, consume small, mesopelagic prey, pre-
sumably one individual at a time (Norris and Dohl
1980). Analyses of stomach contents of spinner dolphins
revealed that they are primarily nocturnal feeders; their
stomachs are full when caught early in the day and
empty when caught at other times of the day (Norris and
Dohl 1980). Stomachs of spinners caught off the coast of
West Africa (Cadenat and Doutre 1959) and in the
eastern tropical Pacific (Perrin et al. 1973) contained
mostly myctophid fish. Fitch and Brownell (1968) found
that two species of myctophid fishes, Benthosema pana-
mense and Lampanyctus parvicauda, accounted for more
436
than 50% of the spinner’s diet in the eastern tropical
Pacific but animals from this area also had large num-
bers of individuals from other mesopelagic fish families
including Gonostomatidae, Melamphaeidae, and Breg-
macerotidae (Fitch and Brownell 1968; Perrin et al.
1973). Mesopelagic squid species, mostly from the
family Enoploteuthidae, also accounted for a large
proportion (nearly 30%) of the prey consumed by
spinner dolphins in the eastern tropical Pacific (Perrin
et al. 1973).
Stomach contents of spinner dolphins caught or
stranded in Hawaiian waters are similar to those caught
in other locations, containing primarily small (2–10 cm)
mesopelagic fish and squid. Norris et al. (1994) found
that more than 50% by number of the contents of
Hawaiian spinner dolphin stomachs were myctophid
fish. Hawaiian spinner dolphins also consumed large
proportions of mesopelagic squid; Abralia trigonura and
A. astrosticta were the most abundant species (Norris
and Dohl 1980; Clarke and Young 1998). One difference
in the diet of Hawaiian spinner dolphins from that of
animals from other locations is the relatively large
contribution of the sergestid crustacean Sergia lucens to
their diet, as well as a small contribution by pasiphaeid
crustaceans (Norris and Dohl 1980; Norris et al. 1994).
Nearly all of the species identified from the stomachs
of Hawaiian spinner dolphins are components of the
mesopelagic boundary community (Reid et al. 1991).
The mesopelagic boundary community is a land-asso-
ciated community that has a composition of fish,
shrimp, and squid that is distinct from the micronek-
tonic community found in waters beyond the archipel-
ago’s slopes (Reid 1994). Many of the species found in
the boundary community migrate diurnally, reaching
depths of 400–700 m during the day and ranging from
the surface to 400 m at night (Reid 1994). The boundary
community also migrates horizontally, moving nearly
2 km toward shore until midnight, when it achieves its
maximum density in waters approximately 1 km from
the shoreline. After midnight, the community then mi-
grates away from shore as it descends (Benoit-Bird et al.
2001). Rather than foraging offshore for the entire night,
spinner dolphins track the horizontal migration of their
prey (Benoit-Bird and Au 2003a). It is hypothesized that
this tracking of their prey allows spinner dolphins to
maximize their foraging time while foraging on the prey
at its highest densities (Benoit-Bird and Au 2003a).
To examine the effect of time and energy constraints
on spinner dolphins, data on the energy content of
spinner dolphin prey and estimates of the foraging needs
of spinner dolphins are presented. Prey energy density
values are crucial inputs to bioenergetic consumption
models but were not available for mesopelagic micro-
nekton in general, or for the specific mesopelagic prey of
spinner dolphins in Hawaii. The effects of various for-
aging costs on energy needs and foraging rates, as well as
the relationship between available prey and spinner
dolphin energy needs, were examined to determine what
factors may be important in driving the diel migration
pattern observed in spinner dolphins. This approach is
conceptually similar to the bioenergetics model of
Stockwell and Johnson (1999), which incorporated for-
aging constraints to predict diel migration observed in
fish. Simple models were also used to predict potential
prey preferences, both by size and by taxonomic iden-
tity. While limited information is available for estimat-
ing the parameters of the model, the relative predictions
of the model are robust to changes in these parameters
and provide information on the importance of prey size
for potential foraging success. Model predictions were
preliminarily compared with results from the stomach
contents of a single stranded animal as well as previously
published gut contents studies.
Materials and methods
Prey value
Animals from the mesopelagic boundary community were captured
alive using an Isaacs–Kidd midwater trawl. Their backscattering
strength (Benoit-Bird and Au 2001), wet weight, dry weight, dis-
placement volume, and caloric content (Benoit-Bird and Au 2002)
were measured. Calorie measurements were made using a bomb
calorimeter.
Total caloric needs
No food intake values are available for spinner dolphins, Stenella
longirostris, either in the wild or in captivity. To estimate the total
energy required for maintenance of a spinner dolphin, the kiloca-
lories per kilogram of body weight required to maintain a captive
striped dolphin, Stenella coeruleoalba (Kastelein et al. 2002a), was
applied to the full range of spinner dolphin weights and compared
with the average maintenance energy values for other delphinids in
captivity.
Stomach contents of a stranded spinner dolphin
As a preliminary test of the viability of the model proposed, the
stomach contents of a stranded animal were examined and com-
pared, along with published reports of spinner dolphin gut con-
tents, with the model predictions. A sub-adult female spinner
dolphin, weighing approximately 45 kg, stranded early in the
morning of 7 July 2002 on Magic Island, Ala Moana Park on
the south shore of Oahu. Despite attempts to save and rehabilitate
the animal, it died. The animal appeared well nourished and no
cause of death was apparent after necropsy. The stomach of the
animal was dissected within 24 h of death, and the identifiable prey
parts, including fish otoliths, squid lenses and beaks, and whole
animals, were frozen, measured, and identified to the lowest taxo-
nomic level possible.
To determine a lower limit to the caloric content of the prey
consumed by this animal, the caloric content was estimated from
the wet weight of the stomach contents. Unfortunately, no catalog
exists for the otoliths of the myctophids or the beaks of the squids
found in the Hawaiian boundary layer so the caloric content of
individual prey items could not be calculated by relating the size
of the hard parts to the length of the prey items. The distribution of
prey caloric content per gram of wet weight was unimodal and not
significantly different from a normal distribution. Because there
were no significant differences between groups, the mean calories
per gram of wet weight for all groups was multiplied by the total
weight of the stomach contents to estimate the total caloric value of
the consumed prey.
 437

The data available on animals from the Hawaiian boundary
community is limited, making it impossible to estimate the length
of prey from the hard parts in the stomach. Two methods were
used to estimate the average length of prey items eaten. First, the
total calorie–length relationship for squid and myctophids was
applied to the estimated average caloric content of each prey item.
Second, we applied the wet weight–length relationship for squid
and myctophids separately to the average wet weight of each prey
item. The total length of any whole animals was directly measured.
Models and parameters
Simple models based on the energetic needs of dolphins and the
energy content of prey were used to predict the number of prey that
an individual spinner dolphin needs to consume. Information on
prey behavior was used to convert these estimates to average rates
of prey consumption. The consequences of foraging costs, derived
from the literature, on these estimates were explored. The effects of
prey size on rate estimates were also examined. The effects of prey
size on foraging estimates allowed prediction of prey size prefer-
ence. Possible preferences for prey type were investigated based on
the acoustic scattering of prey and the swimming ability of prey
obtained from the literature. Finally, information on the distribu-
tion of prey in the field was combined with the foraging models to
understand the limitations faced by spinner dolphins in the wild.
Results
Prey value
The total caloric content of individual prey items from
the boundary community measured with a bomb calo-
rimeter increases with increasing animal length (Fig. 1a).
The relationship can be described by a power function
for myctophid fishes, and a linear function approxi-
mately describes the relationship for shrimp and squid
where E is energy content in calories, and L is total
length in centimeters for all three groups.
EFish ¼ 8.8
L2:64 ; R2 ¼ 0.87; n ¼ 58
EShrimp ¼ 228
L  78.2, R2 ¼ 0.87; n ¼ 14
fit (R2=0.76) after removing the single outlier, a nearly
10-cm-long myctophid. The relationship between the
wet weight of individual animals and their energy con-
tent in calories is shown in Fig. 1b (calories=2,496*wet
weight+55; R2=0.95). The outlier did not significantly
affect this relationship. The distribution of prey caloric
content per gram of wet weight (Fig. 2) was unimodal
and approximated a normal distribution. There were
also no significant differences in the caloric content per
gram of animal wet weight between shrimp, myctophids,
and squid (ANOVA; df=2, P>0.10). The caloric den-
sity of animals from the mesopelagic boundary com-
munity is high (mean 2,837 cal/g wet weight) relative to
food commonly fed to marine mammals in captivity
(Table 1). Only anchovies and sardines, both high in oil
content like mesopelagic animals, come close to the
energy density of the food consumed by spinner
dolphins in the wild (Kizevetter 1971/1973; Honda et al.
1992).
Total caloric needs
The maintenance requirement for a striped dolphin,
54 kcal/kg of dolphin per day (Kastelein et al. 2002a),
compares well with the average maintenance energy
values of 50 kcal/kg of dolphin per day reported for
other delphinids in captivity (J. Pawloski and M. Breese,
personal communication). The estimated kilocalories,
assuming a 54 kcal/kg maintenance need, required for
maintenance each day by spinner dolphins of various
sizes are shown in Table 2. These represent the best
estimates available for the average maintenance energy
needs for a spinner dolphin. Differences in these values
affect the results of the model by shifting curves but do
not change the relative relationships, so for simplicity,
these fixed values are utilized throughout.

ESquid ¼ 214
L  379.6; R2 ¼ 0.80; n ¼ 8 Stomach contents of a stranded spinner dolphin

A single linear relationship can be fitted through The entire contents of the stomach weighed 834 g (wet)
all three groups pooled for simplification of analysis and had a displacement volume of 3.7 l. The identified
(ETotal=388*L)1,139) while retaining a relatively strong contents included 2 intact shrimp (both Sergia fulgens),
Fig. 1 a Prey total caloric
content, measured with a bomb
calorimeter, of mesopelagic
animals from the Hawaiian
mesopelagic boundary layer
as a function of animal length.
b Total caloric value of all three
taxonomic groups of
mesopelagic prey from the
Hawaiian boundary layer was
strongly correlated with wet
weight
438
Fig. 2 Distribution of calories per gram of wet weight for all
mesopelagic boundary community animals measured. No signifi-
cant differences were observed between myctophid fishes, shrimps,
and squids
1,012 fish otoliths (96% were identified as myctophids:
88% Benthosema, 8% Diaphus), 117 whole squids [all
identified as mesopelagic boundary community species
(Reid et al. 1991)], 720 squid eye lenses, and 102 squid
beaks. These prey bodies and parts were interpreted to
represent at least 985 individual prey. Because spinner
dolphins captured in the afternoon typically have com-
pletely empty stomachs (Perrin et al. 1973, 1999), it is
likely that all 985 of the prey items represented in the
stomach contents were eaten the night before the animal
stranded.
The estimated caloric content of the stomach con-
tents was about 2,400 kcal, making the average caloric
content of each prey item about 2,400 cal. Because of
the degree of digestion and consideration of only the
stomach, not the entire gut, this should be regarded as a
minimum estimate of the calories consumed by this
animal. The maintenance energy needs of a spinner
dolphin this size, without considering possible growth
costs, were estimated at 2,440 kcal per day (Table 2).
The size of prey consumed by this dolphin was of
interest for preliminarily examining the results from the
models presented. The total calorie–length relationship
for squid and myctophids was applied to the estimated
average caloric content of each prey item, giving an
Table 1 Mean caloric densities
(calories per gram of wet
weight) for prey items
commonly fed to marine Boundary community
mammals compared with prey Anchovy
of Hawaiian spinner dolphins Sardines
and other midwater Capelin
micronektonic animals Herring
Other midwater vertical
migrators
Mackeral
Cod
Smelt
Squid
estimate of the average squid total length of 11.4 cm and
an average myctophid total length of 7.8 cm. Applying
the wet weight–length relationship for squid and myc-
tophids separately to the average wet weight of each
prey item yielded an estimate for the average squid total
length of 8.4 cm and an average myctophid total length
of 7.7 cm. The total length of the 117 whole squids ex-
tracted from the stomach ranged from 2.4 to 9.6 cm,
with a mean of 7.7 cm (Fig. 3).
Models and parameters
The steps in the derivation of the models and their
parameters build successively; each component of the
model is presented under a separate heading along with
justification for parameter choices, a more detailed
description of the methods used to generate the results,
and the important findings.
Foraging costs
Maintenance energy requirements obtained from captive
studies include only costs associated with feeding and
routine activities, and not those associated with forag-
ing. To understand the importance of food to spinner
dolphin behavior, the effects of foraging costs on energy
needs and food intake rates need to be explored. The
energetic costs experienced by foraging spinner dolphins
come from swimming, diving, acoustic searching, and
‘herding’ of prey and finally from behaviors associated
with the capture of the prey. Spinner dolphins foraging
on the Hawaiian mesopelagic boundary community
move at least 8 km back and forth over the slope of the
island each night while following the horizontal migra-
tion of their prey (Benoit-Bird and Au 2003a). Near the
beginning and end of the prey’s migration, spinner
dolphins probably dive to depths of at least 150 m. Near
the apex of the migration, their diving depths are re-
duced to an average of 25 m (Benoit-Bird and Au
2003a). Hawaiian spinner dolphins feed cooperatively,
in pairs, within relatively large groups (20 or more ani-
mals). They appear to actively concentrate their prey by
swimming in a circle about 25 m in diameter around the
outside edges of the prey (Benoit-Bird and Au 2003a).
Mean caloric density Source
2,837 This article
2,808 Kizevetter 1971/1973
2,300 Honda et al. 1992
1,266–1,863 Martensson et al. 1996
1,480–1,831 Kastelein et al. 2002b; Martensson et al. 1996
1,800 Brooks 1977
1,700 Cox et al. 1996
1,003)1,340 Smith et al. 1997; Martensson et al. 1996
1,200 M. Breese, personal communication
852 Cox et al. 1996

Table 2 Estimated kilocalories required by spinner dolphins each
day for maintenance
Dolphin weight (kg) kcal day)1
Stranded animal 45 2,430
Small adult female 55 2,970
Average adult 65 3,520
Large adult male 75 4,050
Fig. 3 Distribution of squid total length for animals extracted
whole from the stomach of a stranded Hawaiian spinner dolphin
The foraging costs to spinner dolphins of finding
prey, diving to the layer, and ‘herding’ are associated
with prey patches and are relatively isolated from indi-
vidual prey, so they will be referred to collectively as
‘search’ costs. The energetic costs associated with prey
search (swimming, acoustic searching, diving, and
‘herding’) are not likely to be affected by the size of
individual prey. The costs of swimming (see Yazdi et al.
1999) over the island’s slopes to find prey would not be
dependent on prey size. Acoustic prey searching ability
at a large scale is not affected by individual prey size, but
by the distribution of the prey patch or entire scattering
layer because of volume reverberation (Urick 1983).
Costs associated with diving to the prey layer (see Wil-
liams et al. 1999) are probably independent of prey size
because the depth of prey is not strongly correlated with
individual animal size in the upper 150 m (Reid 1994).
Finally, the conserved nature of the observed coopera-
tive ‘herding’ behavior of prey by spinner dolphins
(Benoit-Bird and Au 2003a) suggests relatively constant
effort is used to herd each prey patch. These search costs
are amortized over all prey in each patch successfully
captured in a foraging bout, and consequently, within a
foraging bout they would be the same for each item,
regardless of the item’s caloric content.
Little information is available with which to estimate
search costs for spinner dolphins directly; however, their
effects on spinner dolphin total energetic needs and
foraging rates can be investigated. A range of search
439
Fig. 4 The number of prey items necessary per day as a function of
mean prey length eaten for an average-size spinner dolphin to meet
its basic energy requirements at different fixed search costs per prey
item
costs from 0.00 to 0.15 kcal per prey item was investi-
gated. This represents an increase of 0–8% over the
animal’s maintenance energy during active foraging.
Increases in metabolic rate of 10% for efficient swim-
ming (Yazdi et al. 1999) and 20% or more for diving
have been observed in dolphins (Williams et al. 1999).
Because dolphins may not be swimming and diving for
the entire foraging time, lower costs were used in the
calculations. However, values up to a 50% increase over
maintenance energy do not affect the conclusions of the
analysis. The effect of adding fixed costs on the number
of prey that must be consumed by an average-size
spinner dolphin to meet its maintenance energy costs,
calculated using the estimated net gain of energy for
individual prey of each size, is shown in Fig. 4. In all
cases, the number of prey necessary to meet the animal’s
energy needs decreases with increasing prey size. When
no search cost is assessed and prey average 2.5 cm in
length, a total of nearly 15,000 prey items must be eaten.
When a dolphin consumes large prey, averaging 10 cm
in length (the largest prey available within the boundary
layer), about 825 prey items must be consumed. If search
costs remain the same regardless of prey size, the costs
will have a greater impact on an animal foraging on
small prey. When a meager 0.15 kcal/prey cost is added
(less than the increase in metabolic costs of swimming
for the entire foraging bout), the estimated number of
prey necessary to meet a dolphin’s predicted energy
needs goes up to nearly 40,000 for a 2.5-cm prey item
(Figure 4). This is a more than threefold increase over
the estimate with no search cost. For 10-cm prey, the
estimated increase in the number of prey that must be
consumed when a 0.15 kcal/prey cost is added only
represents a few percent over the number that must be
440
consumed when no cost is assessed. The results can also
be looked at as the estimated total calories that must be
consumed by an individual dolphin as a function of prey
size at different search costs (Fig. 5a). This suggests that
animals should focus foraging effort on prey items about
6 cm in total length or longer to minimize the number of
prey and consequently the total number of calories they
need to consume. Changing the maintenance energy of
the individual or changing the search costs to higher
values has no effect on this prediction.
Search costs are probably independent of prey size,
but prey capture costs probably increase with prey size
because prey mobility also increases. If prey capture
costs increase proportionately with prey size as observed
in fish (Sambilay 1990) and squid (Yatsu et al. 1999), the
simplest case, then the prey capture cost can be ex-
pressed as a percentage of the post-search value of the
prey (net prey value=gross prey caloric content)0.0–
0.15 kcal fixed search cost per prey)capture cost ex-
pressed as a percentage caloric content of each prey;
Fig. 5b). The values selected are within the same range
as the percentage increase in metabolic rates associated
with burst swimming over efficient swimming (5–20%;
Yazdi et al. 1999). This added prey capture cost de-
creases the slope of the total calorie curve between small
and large prey items. Changing the prey capture cost to
a higher value shifts the curve up and further decreases
the difference in the estimated total calories that must be
consumed between small and large prey. However, even
at the lowest fixed search cost (0.05 kcal/prey), it is not
possible to negate the differences between small and
large prey in the estimated total calories that need to be
consumed by the dolphin. Regardless of the prey capture
cost, dolphins are still predicted to preferentially con-
sume larger prey.
Fig. 5 a Total calories necessary per day for an average-size
spinner dolphin to meet its basic energy requirements as a function
of prey size at different fixed per-prey-item search costs. b Total
calories that must be consumed by an average-size spinner dolphin
to meet its basic energy requirements with both a fixed search cost
per prey item and a prey capture cost that increases with prey size
Other costs
Relatively little information is available on the energetic
costs of lactation or pregnancy in cetaceans. Captive
bottlenose dolphins did not change the number of cal-
ories consumed during gestation but increased their
caloric consumption by 52–97% during lactation (Kas-
telein et al. 2002b). Other mammals that feed their off-
spring for a similar amount of time, including hamsters
and humans, also increase their energy consumption by
more than 50% during lactation (Day et al. 2002; Du-
four et al. 2002). Estimated increases in the average-size
female spinner dolphin’s food intake by various per-
centages to represent the energetic costs of lactation over
maintenance energy are shown in Fig. 6. The data in this
figure can also be used to estimate changes in food in-
take due to other causes such as growth, health, and
environmental changes, among others, as these change
the maintenance energy requirements. For example,
growth in a subadult captive dusky dolphin resulted in
approximately a 10% increase in maintenance energy
over the 50 kcal/kg base value (Kastelein et al. 2000),
resulting in a modest increase in estimated total energy
that must be consumed by this growing animal.
How fast must a dolphin eat?
Spinner dolphins have a limited effective foraging depth,
estimated at between 200 and 250 m (Fitch and
Fig. 6 Estimated total caloric needs for the average-size female
spinner dolphin at different non-foraging costs. The costs of
foraging on 7.5-cm-long prey are included at both a fixed, per prey
item level, and a fixed plus percentage foraging cost. The dotted
region shows the range of costs of lactation observed in captive
bottlenose dolphins by Kastelein et al. (2002b). This graph can also
be used to examine the effects of other costs such as growth or
stress. Growth, for example, added a 10% cost to maintenance
needs for a captive dolphin of another species (Kastelein et al.
2000)

Brownell 1968; Dolar et al. 2003) although they may be
able to dive to depths of up to 400 m (Dolar et al. 2003).
This means that the boundary community is inaccessible
to spinner dolphins during the day due to its depth at
between 400 and 700 m at that time. Based on the ver-
tical migration patterns observed in the boundary layer
by Reid (1994) and Benoit-Bird et al. (2001), the
boundary layer is probably accessible to spinner dol-
phins for approximately 11 h of each day. An average-
size adult spinner dolphin eating only the largest prey
available (10 cm in length) would need to average an
estimated 1.25 prey items per minute of its probable
foraging time for maintenance energy. A dolphin con-
suming 2.5 cm prey is predicted to average 22 prey/min
over the 11 h the boundary community is available to
spinner dolphins if no foraging costs are considered.
Adding both prey search and capture costs does not
substantially alter the predicted rate of a dolphin con-
suming large prey, increasing it to only an estimated
maximum of 1.4 prey/min. If the length of prey averaged
7.5 cm, the model estimates that between 3 and 4 prey/
min would need to be consumed, depending on search
and capture costs. If prey were on average only 2.5 cm
long, the estimated number of prey consumed per min-
ute would range between 22 and 46, depending on total
foraging costs.
Possible prey preferences
No strong differences in the amount of energy available
within the different major taxonomic groups of meso-
pelagic prey around Hawaii are apparent, so no
advantage of prey groups is clearly evident (no proxi-
mate analyses were conducted so no conclusions can be
made about other potential differences in food quality).
However, there are probably differences in the costs of
foraging upon the different prey groups, particularly in
prey detection and handling. Assuming spinner dolphins
use echolocation to find their prey, the backscattering
strength of prey items determines the ability of cetaceans
to detect them. Shrimp and squid have a much smaller
backscattering cross-section than myctophids, meaning
that they are weaker echolocation targets (Fig. 7). Al-
though spinner dolphins use a very different sonar signal
than the one used by Benoit-Bird and Au (2001) to
measure scattering from spinner dolphin prey, their
relative scattering strengths probably remain similar.
The difference in scattering strength between fish,
shrimp, and squid has been observed at individual fre-
quencies between 30 and 220 kHz (MacLennan and
Simmonds 1992), covering the entire frequency range of
spinner dolphin echolocation signals (Schotten et al.
2004). Because of these differences in scattering strength,
individual shrimp and squid would be detectable to an
echolocating dolphin over less range than myctophid
fishes. There are also important differences in the ability
of sonar to determine prey size between the prey groups,
significant because of the hypothesized effect of prey size
441
Fig. 7 The relationship between total length of individual prey
taxonomic groups and their dorsal-aspect acoustic backscattering
cross-section at 200 kHz (Redrawn from Benoit-Bird and Au 2001)
on a spinner dolphin’s ability to meet its energetic needs.
While all groups of prey exhibit an increase in back-
scattering strength with increasing length, the differences
in backscatter from shrimp and squid over the range of
available prey sizes is small (Benoit-Bird and Au 2001).
Because backscattering strength is also affected by ori-
entation of the animal (Benoit-Bird and Au 2001) and its
behavior (Arnaya et al. 1989a, 1989b), these small dif-
ferences probably make it difficult for a dolphin to
estimate accurately the size of a shrimp or squid using
echolocation. Myctophid fishes show a much stronger
length–backscattering strength relationship, making it
easier to assess the prey’s size with echolocation before it
is consumed. If echolocation is important in prey
detection and selection in spinner dolphins, myctophids
should be preferred over shrimp or squid of equal size.
Once prey is detected and selected, there are probably
differences in the ability of spinner dolphins to capture
prey of the different groups. Large differences between
groups exist in the swimming rates of individuals of
similar size. Sustained swimming speeds in a species of
shrimp in the same genus, of the same size and body type,
as the most abundant Hawaiian boundary community
shrimp, reach 0.06 m/s, with burst speeds of 0.2 m/s
(Cowles 2001). Similar burst speeds have been observed
in krill, similar in size and body plan to mesopelagic
boundary shrimp, when moving forward; however, re-
verse thrusts can reach 1.1 m/s and can cover 0.6 m (Kils
1979). Hawaiian boundary community myctophids reach
much higher sustained speeds of 0.75 m/s (Reid 1994).
Burst swimming speeds of fish with the size and fin type
of myctophids are estimated at around 3 m/s (Sambilay
1990). Burst speeds of squid (corrected for body length)
are even higher at 8 m/s, with cruising speeds of 0.8 m/s
(Yatsu et al. 1999). Based on swimming speed estimates,
shrimp are probably the easiest common prey for spinner
dolphins to capture, with myctophids intermediate, and
squid the most difficult.
442
The distribution of mesopelagic boundary commu-
nity species and their relative abundance is currently
unknown. However, the combination of ease of detec-
tion and size determination of myctophids with echo-
location and their moderate swimming abilities
compared with the other prey choices suggest that they
should be preferentially consumed over the other two
prey groups. Shrimp and squid would both be more
difficult to detect; however the order of magnitude
slower burst speeds of shrimp suggest that, given equal
sizes, they should be preferentially consumed over squid.
Information on the distribution of these groups will help
to refine these predictions based on diving costs of dol-
phins and relative availability of the prey groups.
Total prey availability
The density of mesopelagic boundary community ani-
mals around the Hawaiian Islands varies as a function of
time, distance from the shoreline, and location. The mean
density of animals found nearshore during the night is
between 5 and 25 animals/m3. Maximum densities can
reach up to 1,800 animals/m3 (Benoit-Bird et al. 2001).
Areas with spinner dolphins have the highest densities
observed (Benoit-Bird and Au 2003a). Utilizing the
acoustic scattering–caloric content relationships estab-
lished by Benoit-Bird and Au (2002) for mesopelagic
boundary community animals, the minimum available
energy within the layer was estimated. To simplify esti-
mates, only the myctophid relationship was utilized,
giving the most conservatively low estimate of energy
possible. The mean caloric density of the mesopelagic
boundary community is estimated at 83 kcal/m3 with a
maximum of nearly 9,000 kcal/m3. The average caloric
density of areas in which spinner dolphins were found
ranged between 112 and 186 kcal/m3, with each single,
discrete patch of prey containing spinner dolphins having
between 1.65*106 kcal and 5.95*106 kcal (Benoit-Bird
and Au 2003a). These patches in which spinner dolphins
were foraging have enough energy to support the esti-
mated maintenance energy needs of between 510 and
1,830 average-size spinner dolphins if they could exploit
all the prey within the patch. The maximum number of
dolphins observed in a single patch at any one time was
26, much lower than could be supported by the patch.
Large, consistent differences in the numerical density
and available energy of prey were observed between
coastlines (Benoit-Bird and Au 2003b). The density of
animals in the nearshore layer off the leeward coast of
Oahu during the night averaged 1.6 times the density
observed off the leeward coast of Hawaii near
Kealakekua Bay (Benoit-Bird et al. 2001). However, the
total available energy per kilometer surveyed off Hawaii
averaged twice that off the leeward coast of Oahu
because of differences in the vertical extent of the layer.
While prey off Hawaii were found in discrete, relatively
small patches (tens of meters across), the prey off Oahu
were found in extensive layers, covering kilometers.
Because of the constancy of these differences, differences
in the population sizes that could be supported by these
prey layers are predicted. If total available prey energy is
more important in determining spinner dolphin foraging
success, the population density of spinner dolphins off
Hawaii’s leeward coast is predicted to be roughly double
that of the leeward Oahu population. If prey density is
more important for foraging success, the density of
spinner dolphins off the leeward Oahu coast should be
higher. Information on the population densities of
spinner dolphins is not available, but differences in
population sizes between locations may provide insights
into the foraging mechanisms used by spinner dolphins.
Preliminary model assessment
Stomach contents from individual dolphins probably
vary widely, particularly when considering only stranded
animals. The estimate of the caloric value of prey in the
stomach of this animal cannot be considered a perfect
measure of the energy consumed, particularly as the
contents of the remainder of the digestive track were not
saved, but it does represent the minimum this animal
consumed on this night. The stomach contents of this
animal show that it is possible for a wild dolphin to meet
the minimum energy needs proposed, consume the large
number of prey predicted by the model in the time it had
to forage, and to fit the volume of this prey inside its
stomach.
Discussion
The energy available within the mesopelagic boundary
community far exceeds the energy necessary to support
the spinner dolphin population off the leeward coasts of
Oahu and Hawaii where spinner dolphins are most
commonly sighted, and it is very densely distributed
(Benoit-Bird and Au 2003b). Each discrete patch in
which dolphins were found but in which no other
boundary community predators were detected had en-
ough prey to support an order of magnitude or two
more dolphins than were detected in each patch. This
suggests that spinner dolphin foraging is limited by time
and consequently efficiency, rather than by the avail-
ability of food. The diel migration behavior of the
mesopelagic boundary community probably eliminates
the possibility of daytime foraging by spinner dolphins
and may alter the costs of foraging throughout the night
as prey depth and prey density change over the migra-
tion.
Analysis of the effects of including even small incre-
mental foraging costs in the model reveals that foraging
by spinner dolphins on small prey probably increases
energetic requirements dramatically. Changes in the ex-
act parameters used in the model change the absolute
numbers predicted, but not the shape of the curve or the
relative predictions. Because of the curve’s shape, pre-

dictions from these simple foraging cost models,
regardless of the costs used, suggest that spinner dol-
phins should selectively forage on prey longer than
about 6 cm. These predictions match well with the
lengths of prey observed in stomach contents. The esti-
mated total length of myctophids (based on partially
digested remains) was 7.7–7.8 cm. Dolar et al. (2003)
found five species of myctophids with average total
lengths (estimated from standard lengths) of 6.3–
13.1 cm. The mean measured length of whole squid from
the single spinner dolphin’s stomach examined in this
article was 7.7 cm. The estimated total length of the
other squid (based on partially digested remains) was
8.4–11.4 cm. The mean total length of squid (converted
from mantle lengths) found in spinner dolphin stomachs
in Hawaii by Norris and Dohl (1980) was about 7.8 cm
and by Dolar et al. (2003) was 7.6 cm. The distribution
of squid lengths measured in this study shows an inverse
relationship to the size distribution of most boundary
community animals; animals in the boundary commu-
nity have a mean length of about 5 cm with a distribu-
tion skewed toward smaller rather than larger animals
(Reid 1994). This suggests that spinner dolphins are
selectively foraging on larger prey, rather than simply
selecting animal sizes at random from the population.
While the sample size for shrimp is small, both this study
and Norris and Dohl (1980) found that shrimp in
Hawaiian spinner dolphin stomachs were 4.8–5.2 cm in
total length, somewhat smaller than predicted by the
estimated foraging costs. Dolar et al. (2003) found
similarly small shrimp in the stomach contents of spin-
ner dolphins in the eastern tropical Pacific. Perhaps the
slow swimming speed of shrimp compared with other
groups reduces the costs associated with foraging upon
shrimp. Differences between taxonomic groups in the
size distribution of the prey may also be a factor.
No matter what size prey spinner dolphins consume
from the size distribution available, the relatively small
size of all available prey probably means that to meet
their energetic needs, spinner dolphins need to forage
quickly. If spinner dolphins use the entire time the
mesopelagic layer is estimated to be within their foraging
range (Fitch and Brownell 1968), they would need to eat
at an average rate of 1.25 10-cm-long prey/min for
maintenance. The foraging rates estimated for mainte-
nance increase dramatically if smaller prey are consumed,
up to 47 2.5-cm-long prey/min if foraging costs (both
search and capture) are considered. If spinner dolphins
consistently consume prey with the average lengths ob-
served in this and the Norris and Dohl studies (1980), that
is, just under 8 cm in total length, then a dolphin would
need to average 3 to 4 prey/min for the entire 11-h po-
tential foraging time. While it is unlikely that this rate is
constant over time, particularly since the depth of the
boundary layer changes by 100 m or more, it does indi-
cate that spinner dolphins are time limited. This limita-
tion lends support to the idea that spinner dolphins need
to follow both inshore/upward and offshore/downward
migrations of their prey (Benoit-Bird and Au 2003a).
443
The total available energy in the mesopelagic
boundary community is substantial and exceeds the
needs of the population of spinner dolphins by orders of
magnitude. Even an individual prey patch can support
many times more dolphins than are actively foraging
within it. This huge excess of energy suggests that den-
sity of prey rather than the total available biomass
probably determines the density, and consequently the
population size, of spinner dolphins. Changes in density
of prey probably have strong effects on the foraging
efficiency (i.e., prey capture rate) of spinner dolphins.
This is evidenced by the strong positive correlation be-
tween spinner dolphins and areas of high prey density
both vertically and horizontally (Benoit-Bird and Au
2003a). It appears that spinner dolphins actively increase
the density of their prey through group foraging
behavior that decreases the size of prey patches while
increasing their aggregation (Benoit-Bird and Au
2003a). For this behavior to be adaptive, this investment
of time by spinner dolphins to create prey areas with
favorable characteristics must be outweighed by
improvements in foraging efficiency within those prey
areas.
The estimates of foraging costs and needs presented
here are based on limited information. While absolute
predictions about spinner dolphin foraging cannot be
made, the results point the way for future research
testing both the expected results and the important
assumptions. The effects of exploring a range of values
in these calculations provide insight into the constraints
under which spinner dolphins forage. A mammal with
relatively high energetic needs, foraging on small mobile
prey individually, but with a limited time to forage, must
forage efficiently, both in terms of capture rate and
selectivity. The importance of food to spinner dolphins
and the effects of these constraints on the patterns ob-
served in their nocturnal behavior are apparent. Spinner
dolphins must be efficient at exploiting their food
resources to meet their energetic needs.
Acknowledgements Marlee Breese, Paul Nachtigall, and Robert
Braun of the Hawaiian Islands Stranding Response Group pro-
vided information on the stranded animal and extracted and saved
its stomach under Dr. Braun’s Letter of Authorization from the
National Marine Fisheries Service. Iris Fischer, Michiel Schotten,
Marc Lammers, Whitlow Au, and Ariel Rivera-Vicente assisted in
the dissection of the stomach. Timothy Tricas generously per-
mitted me to utilize his laboratory and Ariel Rivera-Vicente made
sure I had everything I needed. Gayla Ivey and Ken Longenecker
assisted with the classification of fish otoliths. Whitlow Au, Paul
Nachtigall, and George Losey provided helpful comments on
earlier drafts of this manuscript. Support for this work was pro-
vided by a Leonida Family Scholarship, the ARCS Foundation,
and the Hawaii Institute of Marine Biology’s Marine Mammal
Research Program. This work was also funded by a grant from
the National Oceanic and Atmospheric Administration, Project
R/FM-7, which is sponsored by the University of Hawaii Sea
Grant College Program, SOEST, under Institutional Grant No.
NA16RG2254 from NOAA Office of Sea Grant, Department of
Commerce. The views expressed herein are those of the author
and do not necessarily reflect the views of NOAA or any of its
subagencies. UNIHI-SEAGRANT-JC-03–01 This is HIMB
contribution 1172.
444
References
Acevedo-Gutierrez A, Croll DA, Tershy BR (2002) High feeding
costs limit dive time in the largest whales. J Exp Biol 205:1747–
1753
Arnaya IN, Sano N, Iida K (1989a) Studies on acoustic target
strength of squid II. Effect of behaviour on averaged dorsal
aspect target strength. Bull Fac Fish Hokkaido Univ 40:83–99
Arnaya IN, Sano N, Iida K (1989b) Studies on acoustic target
strength of squid III. Measurement of the mean target strength
of small live squid. Bull Fac Fish Hokkaido Univ 40:110
Benoit-Bird KJ, Au WWL (2001) Target strength measurements of
animals from the Hawaiian mesopelagic boundary community.
J Acoust Soc Am 110:812–819
Benoit-Bird KJ, Au WWL (2002) Energy: converting from acoustic
to biological resource units. J Acoust Soc Am 111:2070–2075
Benoit-Bird KJ, Au WWL (2003a) Prey dynamics affect foraging
by a pelagic predator (Stenella longirostris) over a range of
spatial and temporal scales. Behav Ecol Sociobiol 53:364–373
Benoit-Bird KJ, Au WWL (2003b) Spatial dynamics of a nearshore
micronekton sound-scattering layer. ICES J Mar Sci 60:899–
913
Benoit-Bird KJ, Au WWL, Brainard RE, Lammers MO (2001)
Diel horizontal migration of the Hawaiian mesopelagic
boundary community observed acoustically. Mar Ecol Prog Ser
217:1–14
Brooks AH (1977) A study of the swimbladders of selected meso-
pelagic fish species. In: Andersen NR, Zahuranec BJ (eds)
Oceanic sound scattering prediction. Plenum Press, New York,
pp 565–590
Bunce A (2001) Prey consumption of Australasian gannets (Morus
serrator) breeding in Port Phillip Bay, southeast Australia, and
potential overlap with commercial fisheries. ICES J Mar Sci
58:904–915
Burke BJ, Rice JA (2002) A linked foraging and bioenergetics
model for southern flounder. Trans Am Fish Soc 131:120–131
Cadenat J, Doutre M (1959) Notes sur les delphinides Ouest-Af-
ricains, 5. Sur un Prodelphinus a long bec capture au large des
cotes du Senegal: Prodelphinus longirostris. Bull Inst Fondam
Afr Noire 21:787–792
Clarke M, Young RE (1998) Description and analysis of cepha-
lopod beaks from stomachs of six species of odontocete ceta-
ceans stranded on Hawaiian shores. J Mar Biol Assoc UK
78:623–641
Cowles DL (2001) Swimming speed and metabolic rate during
routine swimming and simulated diel vertical migration of
Sergestes similis in the laboratory. Pac Sci 55:215–226
Cox M, Gaglione E, Prowten P, Noonan M (1996) Food prefer-
ences communicated via symbol discrimination by a California
sea lion (Zalophus californianus). Aquat Mamm 22:3–10
Day DE, Mintz EM, Bartness TJ (2002) Diet choice exaggerates
food hoarding, intake and pup survival across reproduction.
Physiol Behav 75:143–157
Dolar MLL, Walker WA, Kooyman GL, Perrin WF (2003)
Comparative feeding ecology of spinner dolphins (Stenella
longirostris) and Fraser’s dolphins (Lagenodelphis hosei) in the
Sulu Sea. Mar Mamm Sci 19:1–19
Dufour DL, Reina JC, Spurr GB (2002) Energy intake and
expenditure of free-living, lactating Colombian women in an
urban setting. Eur J Clin Nutr 56:205–213
Fitch JE, Brownell RL Jr (1968) Fish otoliths and their importance
in interpreting feeding habits. J Fish Res Bd Can 25:2561–2574
Gorman ML, Mills MG, Raath JP, Speakman JR (1998) High
hunting costs make African wild dogs vulnerable to klepto-
parasitism by hyaenas. Nature 391:479–481
Honda S, Wada T, Matsubara T (1992) Relationship between food
consumption and growth of Japanese sardine Sardinops mela-
nostictus in the laboratory. Bull Fac Fish Hokkaido Univ
56:1–6
Kastelein RA, Elst CA van der, Tennant HK, Wiepkema PR
(2000) Food consumption and growth of a female dusky dol-
phin (Lagenorhynchus obscurus). Zoo Biol 19:131–142
Kastelein RA, Au WWL, de Haan D (2002a) Underwater audio-
gram of a striped dolphin (Stenella coeruleoalba) measured with
narrow-band frequency-modulated signals. J Acoust Soc Am
112:334–344
Kastelein RA, Vaughan N, Walton S, Wiepkema PR (2002b) Food
intake and body measurements of Atlantic bottlenose dolphins
(Tursiops truncatus) in captivity. Mar Environ Res 53:199–218
Kils U (1979) Swimming speed and escape capacity of Antarctic
krill, Euphausia superba. Rep Mar Res 27:264–266
Kizevetter IV (1973) Chemistry and technology of Pacific fish. The
Israel Program for Scientific Translations (trans). U.S.
Department of Commerce, National Technical Information
Service, Springfield, Va., ‘IT72–50019 (Original work published
1971)
MacLennan DN, Simmonds EJ (1992) Fisheries acoustics. Chap-
man and Hall, New York
Martensson PE, Lager Gotaas AR, Nordoy ES, Blix AS (1996)
Seasonal changes in energy density of prey of northeast Atlantic
seals and whales. Mar Mamm Sci 12:635–640
Norris KS, Dohl TP (1980) Behavior of the Hawaiian spinner
dolphin, Stenella longirostris. Fish Bull 77:821–849
Norris KS, Würsig B, Wells RS, Würsig M (1994) The Hawaiian
spinner dolphin. University of California Press, Berkeley, Calif.
Perrin WF, Warner RR, Fiscus CH, Holts DB (1973) Stomach
contents of porpoise, Stenella spp., and yellowfin tuna, Thunnus
albacares, in mixed-species aggregations. Fish Bull 71:1077–
1092
Perrin WF, Dolar MLL, Robineau D (1999) Spinner dolphins
(Stenella longirostris) of the western Pacific and southeast Asia:
Pelagic and shallow-water forms. Mar Mamm Sci 15:1029–1053
Reid SB (1994) Spatial structure of the mesopelagic fish community
in the Hawaiian boundary region. Ph.D. dissertation, Depart-
ment of Oceanography, University of Hawaii, Honolulu
Reid SB, Hirota J, Young RE, Hallacher LE (1991) Mesopelagic-
boundary community in Hawaii: micronekton at the interface
between neritic and oceanic ecosystems. Mar Biol 109:427–440
Sambilay VC Jr (1990) Interrelationships between swimming speed,
caudal fin aspect ratio and body length of fishes. Fishbyte
83:16–20
Schotten M, Au WWL, Lammers MO, Aubauer R (2004) Echo-
location recordings and localizations of wild spinner dolphins
(Stenella longirostris) and pantropical spotted dolphins (Ste-
nella attenuata) using a four hydrophone array. In: Thomas JA,
Moss C, Vater M (eds) Echolocation in bats and dolphins.
University of Chicago Press, Chicago (in press)
Smith RL, Paul JM, Gillispie J (1997) Energy content of Arctic cod
and saffron cod in the northeastern Chukchi Sea. In: Reynolds
JB (ed) Fish ecology in Arctic North America. American
Fisheries Society, Symposium 19. American Fisheries Society,
Bethesda, Md., pp 319–325
Stockwell JD, Johnson BM (1999) Field evaluation of a bioener-
getics-based foraging model for kokanee (Oncorhynchus nerka).
Can J Fish Aquat Sci 56:140–151
Urick RJ (1983) Principles of underwater sound. McGraw Hill,
New York
Williams TM, Haun JE, Friedl W (1999) The diving physiology of
bottlenose dolphins (Tursiops truncatus) I. Balancing the de-
mands of exercise for energy conservation at depth. J Exp Biol
202:2739–2748
Yatsu A, Yamanaka K, Yamashiro C (1999) Tracking experiments
of the jumbo flying squid, Dosidicus gigas, with an ultrasonic
telemetry system in the eastern Pacific Ocean. Bull Natl Res Inst
Far Seas Fish 36:55–60
Yazdi P, Kilian A, Culik BM (1999) Energy expenditure of
swimming bottlenose dolphins (Tursiops truncatus). Mar Biol
134:601–607