The Behavioral Ecology of Shellfish Gathering in Western

Kiribati, Micronesia 1: Prey Choice

Frank R. Thomas
Published online: 29 November 2006
# Springer Science + Business Media, LLC 2006
Abstract Focusing on contemporary shellfish exploitation
among several atoll communities in Kiribati, Micronesia,
this paper examines the relationship between human
foragers and their invertebrate prey via the prey choice or
diet breadth model derived from optimal foraging theory.
Shellfish, like many other reef organisms, are relatively
sedentary and predictable, but these characteristics make
them susceptible to over-harvesting. The research reveals
that shellfish gatherers are foraging in a manner that
matches the predictions of optimal foraging theory. The
work adds to our understanding of optimal foraging
decisions in atoll settings by critically evaluating the
depiction of atoll dwellers as conservationists.
Key words Prey choice
.
shellfish gathering
.
conservation
.
Kiribati
.
Micronesia
Introduction
Research into the behavioral ecology of food acquisition,
better known as optimal foraging theory (OFT), has
blossomed in recent years. Despite important ethnographic
and archaeological applications of foraging models, there
have been relatively few case studies of fishing strategies,
particularly among tropical intertidal gatherers and fishers
(but see Aswani, 1998; Beckerman, 1983; Begossi, 1995;
Bn and Tewfik, 2001; Bird and Bliege Bird, 1997; De
Boer et al., 2002; Sosis, 2002). This two-part study of
shellfish gathering in a contemporary Pacific atoll setting
aims to test several models derived from OFT. This paper
focuses on the prey choice or diet breadth model. A second
article in this series will examine patch switching, patch
sampling, and risk.
Western Kiribati (formerly the Gilbert Islands) consists
of 16 atolls and table reefs spread over 640 km on both
sides of the equator in the west-central Pacific Ocean
between N330 and S245 and E17230 and E17700.
The area is considered part of Micronesia, a set of diverse
archipelagoes, which includes Nauru, the Marshall, Caroline,
and Mariana Islands.
There may be more than 1,000 species of shellfish in
Kiribati, based on estimates recorded in the neighboring
Marshall Islands to the north. Among the open atolls,
high biomass occurs together with relatively high diversity
due to the influence of the nutrient-rich waters of the
equatorial upwelling area (Paulay, 2001). As a result,
shellfish gathering is an important component of daily
subsistence activities among I-Kiribati (indigenous Gilbert
Islanders), with some species targeted for sale in local
markets.
1
Hum Ecol (2007) 35:179194
DOI 10.1007/s10745-006-9066-5
F. R. Thomas (*)
University of the South Pacific, Pacific Studies Program,
PIAS-DG, Suva, Fiji
e-mail: f.r.thomas@sympatico.ca
e-mail: thomas_fr@usp.ac.fj
1
Compared to most finfish catches large quantities of shellfish are
usually required to prepare a meal for a typical household. In that
respect, patterns of marine resource exploitation in Kiribati correspond
to the Pacific-wide focus on finfish as the major source of protein.
However, shellfish gathering increases during periods of daytime low
tides at new or full moon (spring tides). A significant rise in shellfish
consumption has occurred on South Tarawa, the administrative and
commercial center of Kiribati; a phenomenon linked to changes in
water circulation when causeways were built in the 1960s, encourag-
ing the establishment of certain species and increased fertilization by
sewage-driven nutrients (Paulay, 2001). With growing populations and
internal migrations from the outer islands to the capital, there has been
more demand for cheap, easily gathered resources, such as shellfish.
Prey Choice
Optimal foraging theory postulates that all organisms select
food or prey types that maximize their short-term harvest
rate. An increased availability of food increases fertility and
survivorship. Minimizing time spent foraging allows the
pursuit of other fitness enhancing activities. Consequently
natural selection would favor individuals that are more
efficient foragers (Kaplan and Hill, 1992). The optimal diet
is usually determined by comparing the amount of energy
acquired to energy expended, as well as time required to
search and process (handle) each prey type.
The prey choice or diet breadth model reveals the set of
resources that should be targeted by an efficient forager.
This model characterizes foraging behavior in a fine
grained environment where prey types are encountered at
random (i.e., in the same relative proportion throughout the
foraging area). A prey or resource type, which provides the
highest amount of energy acquired to energy expended per
unit time invested in pursuit and handling should always be
included in the diet on-encounter. It follows that when there
is a general increase in environmental productivity (result-
ing in frequent encounters with high-ranked resources), the
variety of food types falling into the optimal diet should be
more restricted than in situations where the environment is
less productive. Thus the inclusion of a prey type in the diet
should depend only on the availability of high-ranked prey
types, and not on its own availability (Emlen, 1966;
MacArthur and Pianka, 1966). In sum, the optimal diet
constitutes a trade-off between search costs and handling
costs (i.e., prey types are added in descending order until
the return rate per unit foraging time is maximized). A more
rigorous algebraic version of this model was developed by
Charnov and Orians (1973), which states that any prey j is
in the optimal diet set if its net energy return (Ej) per unit
handling time (hj) is greater than, or equal to, the average
return rate (including search time) for all prey types of
higher rank:
Ej=hj
X
Li:Ei
.
X
Li:hi 1
Where Li is the rate at which each prey type is
encountered. To determine the optimal diet, three variables
must be known for each prey type: (1) the average expected
net energy gain after encounter with prey type i; (2) the
handling time with an individual of type i after encounter;
and (3) the rate of encounter with prey type i. It is assumed
that a forager knows, on the basis of past experience, the
mean encounter rate, average energy returns, and handling
costs associated with each prey type. Despite certain
methodological problems (for reviews and summaries of
case studies see Kaplan and Hill, 1992; Smith, 1983), the
prey choice model has shown its utility in structuring
questions and predicting the outcome of a number of
foraging decisions (Bettinger, 1991; Krebs and McCleery,
1984; Maynard Smith, 1978; Pyke et al., 1977; Smith and
Winterhalder, 1992).
I-KIRIBATI Shellfish Patches
In most real life situations, resources are not randomly
distributed. Instead, they are clumped, so that their
distribution is said to be patchy or coarse-grained.
Environments that are markedly discontinuous (heteroge-
neous) consist of a patchwork of different resources, in
contrast to uniform or homogeneous environments contain-
ing similar or well-mixed resources (MacArthur and
Pianka, 1966). Where resources occur in patches, the key
assumption of the prey choice model, which states that
search time is equally shared among all prey types, would
seem inappropriate as the forager perceives resource
concentrations, often knows their location, and can accord-
ingly choose an itinerary, in violation of the random
encounter assumption of the model. While the prey choice
model cannot be applied to a set of prey types drawn from
different patches, within a patch, prey types will be
encountered randomly and the prey choice model is
appropriate. In sum, a forager must decide which set of
patches to forage in and how long to forage in each. These
issues will be addressed in a subsequent paper.
Shellfish gathering remains essentially a spatially
bounded activity. The I-Kiribati have long recognized that
specific areas (patches) of the lagoon and ocean habitats
tended to yield different resources or similar kinds of
resources but varying in density (Table I; Fig. 1). The
classification used in this study closely matches the patch
types recognized by the I-Kiribati.
The nearshore patch is primarily exploited for two shell
taxa: Smooth Beach Clam, Atactodea striata, and Pacific
Asaphis, Asaphis violascens. Women and children who
search for food visit this patch at low tide, digging into the
sand matrix. Being closest to shore, this patch is accessible
for relatively long periods of time, although the use of
beach latrines has limited exploitation near densely popu-
lated areas. The sand flat is the most extensive patch on the
lagoon reef flat. Resources there vary according to water
depth, size of sand particles, wave energy, and sand depth.
Several epibenthic, semi-infaunal, infaunal, and sessile
shellfish inhabit this zone. The sand flat supports the
economically important Burnt-end Ark, Anadara uropigi-
melana, Strawberry Conch, Strombus luhuanus, and Pecti-
nate Venus, Gafrarium pectinatum. The lagoon border of
sand flats may support extensive beds of seagrass along the
low intertidal and subtidal regions. Seagrass beds are
productive grounds for a variety of shellfish and are also
180 Hum Ecol (2007) 35:179194
important in fishing because of the high concentration of
nutrients. Beyond the seagrass, the lagoon slope was best
known for its fishing and commercial exploitation by divers
of A. uropigimelana on Tarawa Atoll. Spear fishing and
Giant Clam (Tridacnidae) gathering may take place along
the leeward reef platform in the vicinity of islets. Divers
often take the smaller species of Giant Clam, Tridacna
maxima, and collect the meat for sale in local markets.
Fishing and limited shellfish gathering also occur on the
ocean side of the atolls. As with the lagoon shore, the ocean
side is bordered by a nearshore patch. The reef flat extends
to the reef crest where Giant Clams and a variety of
gastropods can be found.
Methods
Field Methodology
Investigations were carried out intermittently between 1993
and 1998 and focused on communities living on five atolls:
Abaiang, Tarawa, Maiana, Abemama, and Tabiteuea. Data
on various aspects of shellfish gathering were obtained in
three ways: by participant observation (focal follows),
foraging diaries, and from the Shellfish Gatherer Survey
of the Tarawa Lagoon Project, a major interdisciplinary
environmental study of the lagoon (Abbott and Garcia,
1995). The following analysis is based primarily on
observed, quantitative data.
Data were recorded during 73 foraging expeditions
covering 69 days, for a total of 139.63 hours of direct
observation (286.92 forager-hours). Mean group size was
2.6 foragers (SD=0.9, N=146). A total of 59 different
individuals took part in these activities. Foragers were
generally members of the households where the investigator
resided and their neighbors. In addition, 65 foraging trips
were recorded from interviews over a period of 51 days, for
an estimated total of 88.5 h of foraging effort (161 forager-
hours). Mean group size was 3.1 foragers (SD=0.7, N=148).
A total of 19 different individuals were interviewed. A third
source of information was derived from data sheets made
available by personnel of the Tarawa Lagoon Project. This
information greatly expanded the sample size of foraging
events with 83 foraging trips covering 26 days and
approximately 191 h of foraging time (257 forager-hours).
Mean group size was 1.3 foragers (SD=0.7), involving 112
individuals. Although not originally conducted for measur-
ing foraging input and output, the survey nevertheless
included data to assess foraging efficiency, while providing
for an independent check of the harvesting patterns noted
through direct observation.
Analytical Methodology
Foraging activities were measured with a digital stopwatch.
Obtaining complete timemotion records for central-place
handling often proved difficult because processing activities
could be carried out at any time of the day or night. Similar
difficulties were encountered in attempts to measure costs
of acquiring firewood in preparation for boiling shellfish to
facilitate extraction of the animal. It was decided to exclude
the latter costs, while attempting to sample the various costs
of culling inedible parts of prey items. In the present
analysis, energy is assumed to be the maximized currency.
As will be demonstrated, observations are generally
consistent with the predictions of net energy maximization.
In other words, local shellfish gatherers appear to maximize
their overall returns within this particular food class, as
opposed to total caloric production. Although not addressed
in this study, more accurate assessments of the nutritional
value of shellfish can be achieved by examining the entire
range of I-Kiribati foraging patterns. Suffice it to say that
the bulk of energy requirements are derived from carbohy-
drate-rich foods, such as breadfruit, rice, and flour (Pargeter
et al., 1984).
Timemotion records (Nydon and Thomas, 1989) were
converted into estimates of energy expenditure. Energy
levels (kcal/minute/forager) for different age and sex
classes were derived from published sources (Durnin and
Passmore, 1967; Norgan et al., 1974; Ulijaszek, 1995).
Table I Main Shellfish Patches and Resources Found on a Typical Kiribati Atoll
Patch Resource
Lagoon nearshore Atactodea striata; Asaphis violascens
Sand flat Anadara uropigimelana; Strombus luhuanus; Gafrarium pectinatum
Seagrass Anadara uropigimelana; Strombus luhuanus; Gafrarium pectinatum; Trachycardium angulatum; Timoclea marica
Lagoon slope Anadara uropigimelana
Leeward reef platform Tridacna maxima; Hippopus hippopus; Lambis lambis
Ocean nearshore Atactodea striata; Asaphis violascens; Nerita spp.
Reef flat Tridacna maxima; Hippopus hippopus; Turbo setosus; Vasum turbinellum
Reef crest Turbo setosus
Hum Ecol (2007) 35:179194 181
Activity categories specific to shellfish gathering were
matched to their closest published equivalents, corrected
for tropical basal metabolic rate reductions of 10.3% for
males, 3.8% for females, and 7.4% for subadults (cf. Henry
and Rees, 1991).
The values in Table II refer to the activities of an average
adult male and female weighing 68 kg and 58 kg,
respectively. The average weights are derived from a
sample of 18 males and 40 females from Ontong Java
Atoll in the Solomon Islands aged between 20 and 29 years,
reported by Friedlaender and Rhoads (1987) and applied to
this study. The weight of a reference subadult (10
12 years of age) is 35 kg (FAO/WHO/UNU, 1985, p. 136).
Measurements from this PolynesianMicronesian popu-
lation are used in the absence of local data on weights
2
. In
cases where published values are not available, the activity
Fig. 1 Lagoon and ocean
patches from a section of Tar-
awa Atoll (courtesy South Pa-
cific Applied Geoscience
Commission).
2
Although Ontong Java is commonly classified as one of the
Polynesian Outliers in Melanesia, it was shown to have consider-
able PolynesianMicronesian affinities (Mitchell et al., 1987, p. 51).
182 Hum Ecol (2007) 35:179194
specific expenditure rate for a woman weighing 60 kg, aged
between 18 and 30 years, and a subadult weighing 35 kg,
aged between 10 and 12 years, is estimated at 85% and 71%
of the adult male value, using a 70 kg male reference
individual, aged between 18 and 30 years, and a basal
metabolic rate of 22.4 kcal/kg/day (cf. FAO/WHO/UNU,
1985, pp. 133134, 136; Henry and Rees, 1991). Mean age
for observed males was 22.2 years (SD=10.4, N=19), while
the mean for females was 28.9 years (SD=9.1, N=20). A
mean of 11 years old (SD=4, N=20) was obtained by
combining both male and female subadults under the age
of 18.
Fossil fuel and equipment inputs were excluded for
comparability with extant analyses of foraging efficiency.
3
The energy acquired per harvest was determined by
weighing the total edible portion for each prey type, using a
dial scale in the field (5 g increments) and applying the
appropriate caloric values. In circumstances where it was
not possible to measure meat weights separately, conver-
sion actors (ratio of edible soft parts/total weight), obtained
from various sources, were utilized (cf. Bird, 1996, p. 168;
Salvat, 1972; Yamaguchi et al., 1993). Twenty-four
shellfish prey types locally regarded as edible were
analyzed for proximate composition (Table III). Amino
and fatty acid profiles followed the methods described by
Tamaru et al. (1992), with methionine determined as the
sulfone. Energy was then calculated. All analyses are single
trial, except for Gafrarium pectinatum, where the values
represent the average of two trials.
Average productivity of each prey type was established
by calculating on-encounter profitability per bout (load)
E/h (kcal/min) and overall efficiency E/T (kcal/min), where
T is total foraging time (sum of the time spent searching Ts
and handling Th). The number of observations N listed in
the subsequent tables refers to the number of distinct
foraging bouts where a particular prey type was harvested.
For group foraging (when several foragers contributed to
one bag or basin and proceeds were expected to be shared),
a simplifying assumption was made in that time spent in
search and handling was equally shared among foragers,
particularly in situations where relatively dense resources
occurred, with search and handling following each other in
quick succession. While this assumption may be tenuous
among groups possessing different foraging abilities (e.g.,
adults vs. young children cf. Bird and Bliege Bird,
2000), subadults who participated in shellfish gathering
activities essentially confined themselves to searching and
harvesting easily accessible resources (i.e., epifaunal or
shallow, infaunal taxa), and thus probably tended to match
adult performance in terms of foraging efficiency.
Results
Prey Choice
A foraging bout usually takes place within a single patch
type, for example, a sand flat, seagrass bed, or nearshore
biotope. Most households are concentrated on the lagoon
3
It is methodologically feasible to calculate the caloric value of fossil
fuels, but as argued by Smith (1991), this approach significantly
reduces the validity of correlating adaptive costs and benefits with
energetic ones. While the monetary cost of fossil fuels can be
incorporated into labor time and energy measures, there are method-
ological dilemmas arising from the analysis of mixed economies
where currencies such as money and energy are used, or where
overhead costs such as equipment maintenance must be divided
among different productive activities.
Table II Activity Categories and Energy Expenditure Rates for I-Kiribati Shellfish Foraging Time/Energy Budgets
Activity category Published equivalent Male (kcal/min) Female (kcal/min) Subadult (kcal/min)
Process shellfish Sitting working
1
1.2 1.0 0.9
Gather shellfish Standing working
2
1.4 1.2 1.0
Dig in soft sand Dusting
3
3.0 2.6 2.1
Dig in gravelly sand Clear light bush
4
3.4 2.9 2.4
Underwater gathering Underwater swimming
5
5.4 3.8
Breaking Lambis shell Cut tree
6
6.2 5.3
Underwater swimming with load Walking uphill at own pace at 1,800 m
7
7.2
Pry shellfish Husk coconuts
8
6.2 5.2
1
Ulijaszek, 1995: 38
2
Ulijaszek, 1995: 39
3
Ulijaszek, 1995: 36
4
Norgan et al., 1974: 339
5
Durnin and Passmore, 1967: 70
6
Norgan et al., 1974: 342
7
Ulijaszek, 1995: 39
8
Norgan et al., 1974: 341
Hum Ecol (2007) 35:179194 183
side of the islets and harvest shellfish and other resources
from this habitat. Where seagrass occurs, it becomes the
focus of the most intensive gathering pressure. Women and
children concentrate on resources in the intertidal and
subtidal areas of the lagoon where they can easily locate the
telltale signs of their prey. Lagoon slope and leeward reef
platform diving is an activity carried out by males only. The
higher in-patch returns in lagoons compared to ocean
patches is the reason why lagoonal gathering is of
considerable importance. Although foragers are quite
willing to walk relatively long distances on the exposed
intertidal lagoon flat (often in excess of overland distances
from a central-place to the ocean side), they also have more
opportunity to increase their net energy acquisition rate
across the various lagoon patches as the tide comes in and
they are compelled to move closer to shore to the less
productive patches (but still more productive relative to the
intertidal ocean patches) (Thomas, 2002).
Seagrass and Sand Flat Table IV illustrates average
productivity of observed seagrass resources. As landings
from the Tarawa Lagoon Project indicate, species compo-
Table IV Seagrass Foraging: Profitability and Overall Returns
Prey type Gross kcal Net kcal Encounter rate/min E/h (kcal/min)
1
Rank
Trachycardium angulatum N=1 181 179.7 0.1 179.7 1
Anadara uropigimelanaN=10 141.1 SD=213.6 138.7 SD=210 0.8 SD=1.5 61.8 SD=15.6 2
Strombus luhuanus N=7 721.8 SD=716.8 689.4 SD=684.4 5.4 SD=6.7 19.6 SD=3 3
Gafrarium pectinatum N=3 8.3 SD=0.9 7.5 SD=0.4 0.2 SD=0.1 14.4 SD=1 4
E/T (kcal/min)
2
5.3
SD=3.5
1
Values used to calculate E/h can be found in Thomas (1999).
2
E/T refers to the overall returns for this patch. Details about whether it is optimal to include any item amongst the ranked resource types in
descending E/h beyond the highest-ranking item are discussed throughout the text and in Thomas (1999).
Table III Summary of Biochemical Analyses (g/100 g and kcal/100 g raw, wet edible weight)
Taxon Amino acids Fatty acids Glycogen Kcal
Nerita plicata 21.8 0.548 1.55 98.3
Cymatium muricinum 19.7 0.473 1.91 90.7
Strombus luhuanus 19.5 0.419 0.82 85.0
Polinices melanostomus 18.2 0.400 2.06 84.6
Nerita polita 18.2 0.542 1.62 84.1
Vasum turbinellum 17.3 0.431 1.91 80.7
Lambis lambis 17.6 0.288 1.77 80.1
Pitar prora 16.5 0.563 1.64 77.6
Tridacna maxima 14.5 1.112 1.90 75.6
Atactodea striata 15.6 0.807 1.44 75.4
Gafrarium pectinatum 13.3 1.475 1.64 73.0
Barbatia foliata 14.6 0.600 2.18 72.5
Polices tumidus 14.5 0.331 2.63 71.5
Strombus variabilis 13.9 0.408 1.19 64.0
Timoclea marica 12.7 0.681 1.49 62.9
Turbo setosus 12.7 0.723 1.23 62.2
Oliva miniacea 12.7 0.482 1.46 61.0
Hippopus hippopus 9.6 1.256 1.72 56.6
Anadara uropigimelana 11.9 0.373 1.26 56.0
Tridacna gigas 7.9 0.733 3.38 51.7
Spondylus squamosus 10.3 0.491 0.81 48.9
Trachycardium angulatum 9.5 0.533 0.91 46.4
Quidnipagus palatam 6.9 0.289 2.75 41.2
Asaphis violascens 7.7 0.423 1.16 39.2
184 Hum Ecol (2007) 35:179194
sition in seagrass and sand flats is roughly comparable,
except for overall density, which is higher in the former
(Paulay, 2001). Sand flats are more accessible than
seagrass-covered areas, which are basically subtidal, be-
cause they occupy large sections of the mid- to low
intertidal zone. With the exception of Tarawa Atoll,
seagrass is either absent or distributed in small patches on
the outer islands, so that foragers spend most of their time
in sand flats. Despite their seemingly barren appearance,
sand flats provide both shelter and suitable algal foods for
S. luhuanus in areas with mixed soft sand and hard
substrata, while the mid-intertidal is often the focus of G.
pectinatum harvesting.
Table V summarizes sand flat harvests. More hours of
direct observation (42.17) were recorded than in seagrass
patches (20.73) because more time was spent with house-
holds living adjacent to areas largely devoid of seagrass.
It is useful to examine the implications of incorporating
central-place handling or postharvest processing costs.
Anthropologists have for some time considered the appli-
cation of central-place foraging models (Stephens and
Krebs, 1996, pp. 5460) to determine the way in which a
maximum load will affect the ranking of resources located
at a distance from a home base and to establish under what
circumstances resources should be brought whole or be
processed prior to transport (Barlow and Metcalfe, 1996;
Bettinger et al., 1997; Bird, 1996, 1997; Bird and Bliege
Bird, 1997; Jones and Madsen, 1989; Metcalfe and Barlow,
1992; Thomas, 2002). The issue has potential significance
in the analysis of decision-making in whether or not the
comparative difficulty of handling a prey type once
harvested affects its rank order.
Table VI compares E/h and rank order for seagrass prey
types with and without postharvest processing. Inclusion of
postharvest processing does not alter the ranking. However,
in the sand flat (Table VII), only Trachycardium angula-
tum, Barbatia foliata, and A. uropigimelana have unaltered
rankings.
Whether or not postharvest processing is included, T.
angulatum is the highest ranking prey type, but compared
to A. uropigimelana, its apparent density is low. Although
no quantitative data on density are available, this prey is
apparently harvested in equal proportion to its abundance
Table V Sand Flat Foraging and Overall Returns
Prey type Gross kcal Net kcal Encounter rate/min E/h (kcal/min) Rank
Trachycardim angulatum N=1 7.3 7.2 0.01 180 1
Barbatia foliata N=2 29 SD=25.6 21.1 SD=14.6 0.5 SD=0.6 71.8 SD=89 2
Anadara uropigimelana N=16 172.3 SD=345.5 165.6 SD=329.1 1.2 SD=2.1 56.7 SD=20 3
Pitar prora N=1 1.3 1.26 0.01 31.5 4
Polinices melanostomus N=1 33.8 32.4 0.7 24.4 5
Strombus luhuanus N=13 449 SD=402.8 433.7 SD=390.7 3.3 SD=2.5 21.8 SD=4 6
Gafrarium pectinatum
1
N=1 91.3 88 1.4 16.4 7
Oliva miniacea N=1 24.4 22.4 0.4 10.3 8
Spondylus squamosus N=1 92.9 58.3 0.5 9 9
Polinices tumidus N=2 70.3 SD=118.7 66.8 SD=112.9 0.3 SD=0.5 9 SD=9.4 9
Cymatium muricinum N=2 32.9 SD=36.9 30.4 SD=34.4 0.7 SD=0.8 6.2 SD=1.5 10
Strombus variabilis N=1 2.1 1.9 0.02 5.0 11
Gafrarium pectinatum N=8 317.2 SD=112 131 SD=91.8 27.7 SD=52.4 3.3 SD=3.1 12
Gafrarium pectinatum/Pitar prora N=2 198.8 SD=33 70.1 SD=19.9 13.3 SD=14.1 1.4 SD=1 13
Gafrarium pectinatum/Timoclea marica N=2 158.7 SD=75.3 53.7 SD=24.5 28.2 SD=11.1 1.3 SD=0.1 14
Barbatia foliata
2
90.6 214.8 10.7 4.5 15
E/T (kcal/min)
3.6
SD=3
1
With digging considered search.
2
Experimental harvest.
Table VI Seagrass Foraging: Comparison of Profitability and Prey
Type Rank Order Before and After Postharvest Processing
Prey type E/h without
postharvest
processing
Rank E/h with
postharvest
processing
Rank
Trachycardium
angulatum
360.5 1 179.7 1
Anadara
uropigimelana
124.4
SD=31.2
2 61.8
SD=15.6
2
Strombus
luhuanus
89.2
SD=1.5
3 19.6
SD=3
3
Gafrarium
pectinatum
29.3
SD=2
4 14.4
SD=1
4
Hum Ecol (2007) 35:179194 185
(expected encounter rate) in both patches. T. angulatum was
harvested on only 10% of all foraging trips (N=146
combined observed, interview, and Shellfish Gatherer
Survey samples), and the mean number of specimens
gathered per bout was 4.3 (SD=6.9). A. uropigimelana is
a prey type that is both high ranked (always taken on-
encounter) and relatively abundant (therefore common in
the diet, unlike T. angulatum).
S. luhuanus (ranked fourth and sixth in sand flats before
and after postharvest processing), is an interesting prey type
precisely because of the contrast in handling time and its
implications for selectivity. Moreover, the behavioral
attribute of this gastropod to aggregate violates the models
assumption of random encounter of available prey types:
encountering one item increases the probability of encoun-
tering another. Aggregation appears to be a feature of all
members of the genus Strombus (Catterall and Poiner,
1983). Here we have the equivalent of a patch, which can
influence the profitability of the prey type.
Excluding the firmly attached B. foliata and Spondylus
squamosus, G. pectinatum and other infaunal bivalves are
the lowest ranking prey. In a sense, the rank of G.
pectinatum is seemingly at odds with its position as a
targeted prey. Taken at face value, the fact that it is sought
after may very well give credence to the notion that sheer
abundance alone is a determinant of collecting rate. Closer
examination of this prey types characteristics and
extrinsic factors, however, shows that when it is targeted
or otherwise harvested, foragers are indeed behaving
efficiently.
The clumped or patchy distribution of several infaunal
prey, similar to Strombidae, may be instructive here.
Because of their density, it is not uncommon for a forager
to dig up two or more prey items at once. Second, prey may
lie so close to the surface that they are visible to a forager
walking in either seagrass or sand flat. Field handling is
thus reduced, and is comparable to the costs incurred while
harvesting semi-infaunal or epibenthic prey. Some of the
observed variability in E/h for G. pectinatum can be
attributed to whether the prey was buried or lying near the
surface. The third aspect influencing foraging decisions is
the tidal cycle that repels or attracts efficient foragers to and
from the mid-intertidal where overall returns may be
maximized under a different set of constraints. These
migrations are important for understanding the wider
problem of patch switching. With the exception of two
foraging trips, G. pectinatum was targeted when relatively
high water levels, often accompanied by windy conditions,
would undoubtedly have prevented efficient foraging in the
low intertidal because of reduced visibility. The exceptions
include one observed and one interview case. In both
instances, the presence of small children, who needed to be
supervised while adults collected, probably would have
reduced foraging efficiency in the deeper sections of the
sand flat.
The remaining prey types were not specifically targeted,
either because their overall density precluded a focused
harvest or that environmental conditions effectively limited
the duration of the foraging process, such as for shallow
burrowing Polinices spp. and Oliva miniacea whose
characteristic trails in the sand can be spotted only at
extreme low tide. On-encounter, however, they are fre-
quently taken, because they differ little in terms of field
handling costs compared to the other prey types.
Table VII Sand Flat Foraging: Comparison of Profitability and Prey Type Rank Order Before and After Postharvest Processing
Prey type E/h without postharvest processing Rank E/h with postharvest processing Rank
Trachycardium angulatum 361.1 1 180 1
Barbatia foliata 140.3 SD=184.3 2 71.8 SD=89 2
Anadara uropigimelana 121.6 SD=30.3 3 56.7 SD=20 3
Strombus luhuanus 94.1 SD=16.2 4 21.8 SD=4 6
Oliva miniacea 80.2 5 10.3 8
Pitar prora 64.1 6 31.5 4
Polinices melanostomus 58 7 24.4 5
Strombus variabilis 52.2 8 5 11
Cymatium muricinum 49.8 SD=11.7 9 6.2 SD=1.5 10
Polinices tumidus 43 SD=37.7 10 9 SD=9.4 9
Gafrarium pectinatum
1
31.1 11 16.4 7
Spondylus squamosus 9.3 12 9 9
Gafrarium pectinatum 4.6 SD=5.1 13 3.3 SD=3.1 12
Gafrarium pectinatum/Pitar prora 1.8 SD=1.2 14 1.4 SD=1 13
Gafrarium pectinatum/Timoclea marica 1.6 SD=0.1 15 1.2 SD=0.1 14
Barbatia foliata
2
4.5 16 4.5 15
1
With digging considered search
2
Experimental harvest.
186 Hum Ecol (2007) 35:179194
Prey that are avoided on-encounter include B. foliata and
S. squamosus, unless they are detached from their matrix or
weakly fastened. As in the case for the occasional infaunal
bivalve lying close to the surface, a weakly attached bivalve
may be considered a different prey type than one firmly
embedded even if they are the same taxon. This encounter
variability is an example of partial preferences, and
underscores the importance of context when analyzing prey
choice (Stephens, 1985). Given the fact that foragers
seldom venture into the lagoon with a cutting implement,
some opportunities to increase E/T are probably missed
when one encounters weakly attached prey.
In most observed cases and the assessment of other data
sources, the addition of prey types increased overall returns.
Apparent departure from optimal foraging was noted in
nine cases, including six observations. All involved the
gathering of infaunal prey (mostly G. pectinatum) from the
mid-intertidal when foragers decided to move closer to
shore with the rising tide. But as previously noted, this
behavior is suboptimal only when compared to favorable
gathering conditions in the low intertidal and shallow
subtidal areas of the lagoon, and a sufficiently high
encounter rate with high ranking prey. To some extent, this
resembles patch switching, even if the sand flat was defined
as a single patch type. When these two sets of activities are
combined (i.e., searching and harvesting S. luhuanus and A.
uropigimelana lagoonward, and digging in the mid-
intertidal), E/T is depressed. However, because of changed
constraints linked to the rising tide and its attendant
influence on foraging efficiency and possible risk, foragers
are in fact maximizing their returns. Although one cannot
say that foraging in the mid intertidal is in strict violation of
the simultaneous search assumption (there is still the
possibility of encountering other prey types, including the
occasional A. uropigimelana and S. luhuanus), an efficient
forager should harvest infaunal bivalves and forego the
benefits of searching and handling other prey. Therefore,
under different sets of constraints as reported here, the fine
grained assumption is only weakly supported. Another
example of changed constraints resulted in what appeared
to be a further illustration of suboptimal harvest. In this
instance, foragers left the low intertidal because of
approaching nightfall, which would have hampered the
search for both epibenthic and semi-infaunal prey, and
delayed the return home. Harvesting continued closer to
shore by digging for G. pectinatum.
There were several cases from the Tarawa Lagoon
Project seemingly at odds with the idea of foraging
efficiency. Although the quality of the data prevented a
better assessment, conditions analogous to those described
above are suggested, such as harvesting G. pectinatum from
seagrass, possibly because they were visible from the
surface, or gathering S. squamosus that might have been
detached from their matrix.
Table IX Nearshore Foraging: Comparison of Profitability and Prey Type Rank Order Before and After Postharvest Processing
Prey type E/h without postharvest processing Rank E/h with postharvest processing Rank
Nerita plicata
1
21.8 1 2.6 1
Asaphis violascens 5 SD=7.9 2 1.3 SD=2.3 2
Nerita polita 1.9 SD=1.1 3 0.7 SD=0.6 3
Atactodea striata 1.2 SD=0.6 4 0.6 SD=0.4 4
Asaphis violascens/Quidnipagus palatam 0.9 SD=0.1 5 0.7 SD=0,04 3
Quidnipagus palatam
2
0.2 SD=1.2 6 0.4 SD=1.2 5
1
Experimental harvest.
2
Experimental harvest.
Table VIII Nearshore Foraging: Profitability and Overall Returns
Prey type Gross kcal Net kcal Encounter rate/min E/h (kcal/min) Rank
Nerita plicata
1
N=1 126.3 101.1 45.7 2.6 1
Asaphis violascens N=9 263.3 SD=115.1 81 15.1 SD=20.6 1.3 SD=2.3 2
Asaphis violascens/Quidnipagus
palatam N=2
178.8 SD=214 145.6 SD=44.6 7.6 SD=4.5 0.7 SD=0.04 3
Nerita polita N=2 172.4 SD=77.4 74.5 SD=73.5 64.8 SD=35.3 0.7 SD=0.6 3
Atactodea striata N=2 118.4 SD=60.8 21 SD=4.6 272.8 SD=179.9 0.6 SD=0.4 4
Quidnipagus palatam
2
N=3 191.4 SD=201 18.3 SD=189.2 73.3 SD=73.3 0.4 SD=1.2 5
E/T (kcal/min)
0.6
SD=0.5
1
Experimental harvest.
2
Experimental harvest.
Hum Ecol (2007) 35:179194 187
Lagoon and Ocean Nearshore Unlike seagrass and sand
flats, nearshore patches on both lagoon and ocean sides
support a limited variety of prey types, although densities
may be locally abundant. Because of the restricted number
of observations, returns for A. violascens (the most often
targeted nearshore prey type) from both lagoon and ocean
patches were combined, because they are essentially
similar. Table VIII shows the results of observed nearshore
harvests, while Table IX compares profitability and prey
type rank order with and without postharvest processing.
While A. violascens is the primary focus of nearshore
harvesting, it is probably not by virtue of being more
abundant than other nearshore prey types, such as the
smaller Nerita plicata or A. striata. In fact, on the basis of
the inverse correlation between abundance and meat yield
(McNab, 1963), smaller prey are expected to be more
abundant. Consequently, there is compelling reason to
believe that A. violascens is gathered because of its
profitability. Of course, new constraints, such as increasing
levels of nearshore pollution associated with the use of
public latrines, may be determining factors in foraging
decisions
4
A. striata can be a focal prey, but nearly always
for a segment of the human population (i.e., infants who
can easily digest small bivalves). The constraint here lies in
the infants ability to digest larger prey, so that a forager
may decide to focus on A. striata even if it means that
overall returns could be depressed.
Most foraging bouts within nearshore patches (N=40,
combined samples) involved the harvesting of a single prey
type, mainly A. violascens. In the few cases where more
than one prey was included in a catch (N=7, five
observations and two interviews), the addition of lower
ranking prey increased overall returns, except in one instance
where the focus shifted from gathering A. violascens to A.
striata. Not surprisingly, the adult male was planning to
secure a meal for his infant son. Otherwise, he may very
well have continued to forage for A. violascens.
Lagoon Slope Over a decade ago, the lagoon slope on
South Tarawa became the focus of an intensive A.
uropigimelana fishery. It is possible that these offshore
beds were new and made more attractive by declining fish
stocks, which prompted commercial fishermen to exploit
the largely undisturbed A. uropigimelana shellfish beds.
The year 1993 coincided with the rapid expansion of this
fishery. Two years later, however, few divers were seen. A
number of informants claimed that because of a tuna
surplus, the price of fish had dropped, thus contributing to a
decline in shellfish harvesting. Other informants stated that
lagoon slope gathering was no longer profitable because of
stock declines, presumably as a result of overharvesting.
Table X shows profitability and overall returns for this
patch. Because all divers were men (who do not generally
process their catch after harvesting) E/h only reflects field
handling. At any rate, a prey destined for sale, as was the
case for most shellfish gathered from the lagoon slope,
obviously frees foragers from postharvest processing costs.
From the consumers point of view, T. angulatum and A.
uropigimelana obviously provide large meat packages and
minimal processing compared to S. luhuanus.
In 80% of the cases from South Tarawa (N=10), divers
specifically targeted A. uropigimelana for sale. The two
remaining divers collected for subsistence, and on both
occasions, gathered the lower ranking S. luhuanus as well.
In both cases, however, E/T was raised.
While overall returns exceed those calculated for
seagrass and sand flats, major constraints facing foragers
include the need to secure a canoe or a floating device to
travel to the patch and hold their catch, as well as the
strenuous nature of the work: divers commonly free dive to
36 m for many hours each day to gather marketable
shellfish. A less obvious constraint may relate to gender
roles. Shellfish gathering is primarily a female activity, but
4
Although N. plicata is the highest ranked resource in the nearshore
patches, it may be avoided to a greater extent than A. violascens
because of its proximity to the surface and evidence of pollution. This
highlights the type of patch classification used in this study, which
could benefit from greater refinements to take into account horizontal
and vertical differences among various benthic biotopes with variable
geomorphology and, consequently, species assemblages. As it stands,
the present classification can be said to violate random cropping
(Anderson, 1979; Sutherland, 1982) for the above resources, as they
are not strictly handled simultaneously. Because N. plicata occurs at
shallower depths than A. violascens, a forager is faced with the
decision to either handle the former by simply gathering this prey
type from the surface or expend greater effort by digging into the
gravelly matrix where the latter are concentrated.
Table X Offshore Foraging: Profitability and Overall Returns
Prey type Gross kcal Net kcal Encounter rate E/h (kcal/min) Rank
Trachycardium angulatum N=2 19.9 SD=17.8 19.7 SD=17.7 0.03 SD=0.02 446.9 SD=126.5 1
Anadara uropigimelana N=10 2,257.1 SD=1,028.4 2,206.4 SD=1,1005.3 2.7 SD=1.3 217.9 SD=0.7 2
Strombus luhuanus N=2 877.7 SD=598.1 829.4 SD=565.2 23.5 SD=31.3 86 SD=0 3
E/T (kcal/min)
15.4
SD=9.6
188 Hum Ecol (2007) 35:179194
it is considered improper for women to dive, and so they
must restrict their work to intertidal and shallow subtidal
patches.
Leeward Reef (Diving) When the I-Kiribati from the more
populated windward islets venture out to the leeward reef
and adjacent islets, they often target T. maxima. Additional
prey encountered include Hippopus hippopus and Lambis
lambis, whose profitability is illustrated in Table XI.
In calculating E/T, the costs of search and handling non-
shellfish resources were also incorporated
5
. Profitability
and corresponding rank for each prey type before and after
postharvest processing are shown in Table XII.
The addition of prey types, including fish and octopus,
among the observed foraging bouts raised overall returns,
except in one instance where E/h for T. maxima (16.6 kcal/
min) the lowest ranking prey reduced E/T from 29.6 to
27.6 kcal/min. However, because the primary motivation
was to collect for income, one cannot say that divers were
foraging suboptimally
6
.
Leeward reef (Walking) Many leeward reef islets currently
do not support permanent settlements. However, some are
temporarily occupied by fishermen who may process fish,
holothurians (seaslugs), and T. maxima. Although most
shellfish resources gathered from the leeward reef require
simple diving gear and a canoe, foraging on foot at low tide
in the intertidal zone at appropriate localities may yield a
similar range of prey types, including T. maxima, H.
hippopus, and L. lambis.
In the process of searching for resources at low tide,
foragers often encounter H. hippopus distributed in what
appeared to be two distinct patterns: the bivalves occur
either singly or in clumps. In the latter case, clumping is
sometimes associated with fish traps, suggesting the
presence of Giant Clam gardens (Thomas, 2001b).
Consequently, the relative high density of H. hippopus is
sometimes explained by deliberate introductions, with
humans responsible for the distribution of this particular
resource. Return rates for pedestrian leeward reef foraging
are presented in Tables XIII and XIV.
Energy expenditure rates and field handling time differ
from leeward reef diving. Because the reef platform
between islets is searched at low tide by walking, gathering
is comparable to standing working (see Table II). In all
instances, at least one group member carried a knife in
anticipation of locating Giant Clams. When more than one
prey type was harvested, overall returns increased for all
observed cases.
Deeply embedded T. maxima (usually small specimens)
were often bypassed because foragers judged them too
Table XI Leeward Reef Foraging (Diving): Profitability and Overall Returns
Prey type Gross kcal Net kcal Encounter rate/min E/h (kcal/min) Rank
Hippopus hippopus N=4 494.3 SD=364.3 482 SD=361.2 0.01 SD=0.01 152.2 SD=88.4 1
Lambis lambis N=5 1,036 SD=1,338.2 1,015 SD=1,318 0.1 SD=0.1 85.1 SD=34.8 2
Tridacna gigas N=2 898.3 SD=747.6 859.6 SD=736.5 0.01 SD=0.04 55.5 SD=78 3
Tridacna maxima N=6 586.2 SD=452.2 532.9 SD=410.7 0.3 SD=0.2 27.5 SD=26.1 4
E/T (kcal/min)
16.3
SD=11.5
6
Money and energy currencies are kept distinct in this analysis. It is
sometimes difficult to separate artisanal fisheries into commercial and
subsistence operations (Adams et al., 1999). The FAO (1998) reports
that the nearshore commercial fish catch in Kiribati is principally
made up of reef- and deep-slope fish (54%), shellfish (25%), and
pelagic species (21%).
5
Because shellfish gathering is not the primary focus of fishing
activities in the leeward reef patch (except for T. maxima), all non-
shellfish prey types needed to be taken into account in determining
overall efficiency. Published data (Leung et al., 1972; Murai et al.,
1958; Sidwell et al., 1974) were used for gross caloric values, and the
categories underwater swimming and sitting working from
Table II were applied to calculate energy expenditure associated with
non-shellfish resources, including octopus, eel, and several varieties of
reef fish. Table XI lists only the Latin binomials for the shellfish prey
types.
Table XII Leeward Reef Foraging (Diving): Comparison of Profit-
ability and Prey Type Rank Order Before and After Postharvest
Processing
Prey type E/h without
postharvest
processing
Rank E/h with
postharvest
processing
Rank
Lambis
lambis
525.7 SD=254.6 1 85.1 SD=34.8 2
Hippopus
hippopus
287.2 SD=136.6 2 152.2 SD=88.4 1
Tridacna
gigas
169.2 SD=119.6 3 55.5 SD=78 3
Tridacna
maxima
64 SD=69.3 4 27.5 SD=26.1 4
Tridacna
gigas
1
12.5 5
1
Experimental harvest (failed pursuit).
Hum Ecol (2007) 35:179194 189
difficult to extract. Up to eight H. hippopus concentrated
inside a fish trap were left in place, as these were
considered off limits because fish traps and resources found
adjacent to them were said to belong to an individual or
extended family. However, a Tridacna gigas specimen,
located about 20 m from the shoreline, was promptly
harvested. Although it is clear that it had been carried to
this location, the absence of a fish trap meant that it could
be collected in accordance with the open access regime
governing nearshore environments and their resources.
Reef Flat Compared to the nearshore ocean patch, which is
sometimes visited when tides or weather prevent efficient
foraging elsewhere, the reef flat does not seem to be the
focus of much shellfish gathering, even during the most
favorable low tides. On occasion, one sees small groups of
fishermen casting their line, but the windward section
remains largely devoid of human subsistence activities. A
notable exception is the presence of fish traps. Neverthe-
less, the reef flat is recognized as a distinct patch where T.
maxima or H. hippopus may be gathered if they are
encountered in the course of fishing trips. There are also
several other shellfish resources that are considered edible.
Results of foraging efforts are presented in Tables XV and
XVI.
The addition of lower ranked prey increased overall
returns among all observed bouts. The experimental
handling of T. maxima was the notable exception: in 48.9
minutes of search, one female forager encountered 0.02 H.
hippopus and 0.1 T. maxima/min. Net energy per encounter
was 80.5 kcal and 0.3 kcal for each prey type, while
handling time per encounter was 1.41 min and 2.88 min.
Profitability for H. hippopus was 57.1 kcal/min, which
resulted in an overall rate of 1.6 kcal/min. Because
profitability for T. maxima was 2 kcal/min (E/h<E/T), it
followed that overall returns dropped to 1.2 kcal/min with
the addition of this lower ranked prey.
Reef Crest Observations and interviews showed limited use
of the reef crest. As a windward ocean patch, it appears
even less significant than the reef flat. Although the sample
for experimental harvests is small (only two observations),
overall returns for the gastropod Turbo setosus are low
(Mean=0.2 kcal/min/load, SD=0.1), which is less than the
average for the combined lagoon and ocean nearshore
patches (0.6 kcal/min).
Efficient foraging can only be accomplished during
extreme diurnal low tides. Foragers must first walk across
the reef flat and then usually progress very slowly along the
reef crest in search of prey. The window of opportunity is
brief. As soon as the tide rises, search becomes more
difficult and sometimes hazardous, as foragers take precau-
tionary measures lest they are knocked off their feet and
injure themselves against the sharp coral. Productivity may
be higher around the time of the full moon. According to
local sources, T. setosus migrates to the coral surface during
that period and is thus easier to locate.
The reef crest supports a variety of gastropods well
adapted to a high-energy environment. Drupa spp., Conus
spp., and Trochus sp. were encountered, in addition to T.
setosus. However, only the latter was regarded as edible.
Returns for the single prey type are presented in
Table XVII. Profitability includes postharvest processing.
T. setosus is the largest resident gastropod. Overall returns
could be substantially higher around the full moon, but this
would require quantitative data. However, even under the
Table XIII Leeward Reef Foraging (Walking): Profitability and Overall Returns
Prey type Gross kcal Net kcal Encounter rate/min E/h (kcal/min) Rank
Lambis lambis N=4 957.8 SD=908.6 939.4 SD=890.2 0.1 SD=0.1 283.1 SD=142.2 1
Tridacna gigas
1
N=1 672.1 668.7 0.01 271 2
Hippopus hippopus N=5 1,379.9 SD=1,009 1,367.5 SD=1,002.7 0.04 SD=0.04 131.9 SD=52.1 3
Tridacna maxima N=3 409.5 SD=361.7 392.7 SD=351.6 0.1 SD=0.01 28.6 SD=10.5 4
E/T (kcal/min)
7.7
SD=3.5
1
Processed entirely in the field.
Table XIV Leeward Reef Foraging (Walking): Comparison of Profitability and Prey Type Rank Order Before and After Postharvest Processing
Prey type E/h without postharvest processing Rank E/h with postharvest processing Rank
Lambis lambis 2,697.6 SD=1,141.9 1 283.1 SD=142.2 1
Hippopus hippopus 366 SD=247.3 2 131.9 SD=52.1 2
Tridacna gigas 271 3
Tridacna maxima 86.7 SD=81.7 4 28.6 SD=10.5 3
190 Hum Ecol (2007) 35:179194
most favorable conditions, returns from the reef crest would
still remain below the average for the other patches,
especially since T. setosus is a vulnerable species (Villiers
and Sire, 1985). Repeated forays, accompanied by brief, but
intensive harvesting could drastically reduce the number of
T. setosus, thus making reef crest foraging a relatively
unproductive venture.
Implications for Conservation
The prey choice model offers a simple way to assess the
hypothesis of short-term costs and gains in food acquisi-
tion, assumed by OFT, and to help better understand
alleged conservation practices by contemporary indigenous
societies. As illustrated by Alvard (1993; 1994; 1995) in
work among an Amerindian community in Peru, a
behavioral ecological approach provides the framework to
distinguish between conservation behavior per se from its
effects, the latter defined as epiphenomenal conservation.
It follows that conservation can be seen as accepting short-
term costs for long-term gains. Whereas other criteria can
be used (Ruttan and Borgerhoff Mulder, 1999; Smith,
1995), this definition of conservation is sufficiently
rigorous to assess deviation from short-term harvest rate
maximization. While conservation of natural resources
among I-Kiribati might take place in other contexts subject
to testing, this case study of shellfish gathering has
demonstrated that foragers are behaving according to the
predictions of the prey choice model.
The previous discussion of foraging decisions by
patches, including instances where prey types were
avoided, supports the idea that foragers are seeking to
maximize their short-term energy gains by adding prey
types to increase overall return rates. High ranking prey
such as A. uropigimelana are always taken on-encounter.
Moreover, there seems to be no size/age discrimination in
harvesting within prey categories: smaller/subadult speci-
mens of S. luhuanus, for instance, are harvested to the same
degree as mature individuals. Such behavior is inconsistent
with a conservation strategy. For S. luhuanus at least, it
clearly does not pay to refrain from taking juveniles with
the aim of reaping benefits in the future because these
gastropods are highly mobile, and therefore, an opportunity
lost today is probably lost forever, even though foragers can
be generally assured of abundant supplies of this resilient
type.
A prey which is seldom or never considered a focal or
targeted resource in no way implies that it is given special
Table XV Reef Flat Foraging: Profitability and Overall Returns
Prey type Gross kcal Net kcal Encounter rate/min E/h (kcal/min) Rank
Spondylus squamosus N=1 24.5 24.3 0.1 121.3 1
Tridacna maxima N=2 119.1 SD=152.4 116.7 SD=150.5 0.02 SD=0.10 43.7 SD=44.2 2
Hippopus hippopus N=3 185.2 SD=237.3 177.1 SD=228.6 0.05 SD=0.1 28.9 SD=27 3
Turbo setosus N=1 9.3 8.6 0.1 10.1 4
Strombus variabilis N=2 141.4 SD=162.1 127.2 SD=149.2 1.6 SD=2 4.9 SD=0.4 5
Vasum turbinellum N=4 72.6 SD=69.2 47.3 SD=49.5 1.7 SD=2 1 SD=0.3 6
Tridacna maxima
1
N=1 79.4 40.6 0.1 2 7
E/T (kcal/min)
1.2
SD=1.2
1
Experimental harvest.
Table XVI Reef Flat Foraging: Comparison of Profitability and Prey Type Rank Order Before and After Postharvest Processing
Prey type E/h without postharvest processing Rank E/h with postharvest processing Rank
Hippopus hippopus 2,269.1 SD=1,839.7 1 28.9 SD=27 3
Spondylus squamosus 243.4 2 121.3 1
Turbo setosus 115.5 3 10.1 4
Tridacna maxima 107 SD=103.2 4 43.7 SD=44.2 2
Strombus variabilis 52.8 SD=0.1 5 4.9 SD=0.4 5
Vasum turbinellum 28.5 SD=1.7 6 1 SD=0.3 6
Tridacna maxima
1
1.9 7 2 7
1
Experimental harvest.
Hum Ecol (2007) 35:179194 191
consideration, for instance to assist in conserving existing
stocks for some future reward. Thus, the high ranked T.
angulatum, occurring in both seagrass and sand flat
patches, is not mentioned as a focal prey (unlike A.
uropigimelana) because it is relatively uncommon, not
because it should be avoided. From what could be
observed, the relative rarity of this prey type in the harvests
does not appear to be linked to the need to restrict
gathering: T. angulatum is the highest ranking prey in both
lagoon patches and is thus predicted to always be harvested
on-encounter.
The case for S. squamosus and B. foliata underscores the
concept of encounter variability and partial preferences.
Firmly attached S. squamosus and B. foliata have different
return rates than detached or weakly attached specimens. If
a forager bypasses prey types in the former state, the
behavior should be entirely consistent with return rate
maximization. Moreover, the lack of an appropriate extrac-
tive technology, such as a knife or crowbar, is a constraint
that limits the foragers ability to efficiently harvest a
resource, which would otherwise increase overall returns.
Knowing the full range of constraints imposed upon
foraging decisions is further illustrated by nearshore
resources: A. violascens, Nerita spp., and A. striata are
sometimes avoided to prevent pathogenic transmission.
Technological constraints, together with unfavorable tidal
conditions, could prevent specimens of Giant Clams from
being efficiently harvested. Given that T. maxima is a
vulnerable prey type (Alcazar and Solis, 1986), how are we
to interpret occasional avoidance? Again, foraging behavior
shows that small, deeply embedded specimens are
bypassed, while more easily accessible bivalves are
harvested. The case for H. hippopus discovered in the
vicinity of a fish trap poses an interesting problem, because
it did appear to involve a short-term cost, but not in the
sense of hoping to derive a long-term gain from the
resource. The bivalves had been placed next to a fish trap,
suggesting they were off limits to outsiders. In other
circumstances involving transported Giant Clams, T. gigas
and H. hippopus, pilferage was common. However, because
foragers in the above instance were searching on the reef in
broad daylight in an open setting where fishermen could
monitor each other, the foraging party decided to leave the
shells in place for fear of retribution.
Atolls are often described as marginal habitats for human
existence, as reflected in the size of habitable landmass and
in the distribution of resources. Compared to terrestrial
ecosystems, the lagoon and reef environments surrounding
the islets provide a wide range of resources. The key to this
high level of productivity is the rapid internal recycling of
nutrients within the ecosystem (Marsh, 1987). However,
even atoll marine environments do not match the level of
productivity found near continental margins or around most
volcanic high islands (Wiens, 1962). Intuitively, therefore,
several authors have suggested that because of these factors
atoll societies were more keenly aware of resource
limitations and, as a result, devised various conservation
strategies. In effect, they would epitomize, at least prior to
Western contact, the idea of the ecologically noble savage
(Redford, 1991). While debate over the effectiveness of
alleged precontact conservation strategies will likely con-
tinue, contemporary observations may assist in critically
evaluating some long-held beliefs and assumptions about
the relationship between humans and their environment on
small, resource poor islands or in other marginal habitats.
Indigenous societies undoubtedly possess a vast knowledge
of their respective environments, which often rivals
Western scientific understanding. However, such knowl-
edge does not necessarily entail a concern for the
environment beyond what it can provide for the immediate
benefit of those extracting resources (Healey, 1993;
Pernetta and Hill, 1984). Moreover, it remains to be
demonstrated whether such knowledge is linked to a
conservation ethic or a user-oriented conservation strat-
egy (McNeely et al., 1995; Ruddle et al., 1992).
7
As Dwyer
(1994) argued, the modern conservation movement is often
at odds with the social and economic reality facing many
indigenous peoples, and as the local political elite often
relies on exclusionary rules that prevent segments of a
population from accessing resources in an increasingly
crowded world. In effect, such communal systems tend to
conserve, but only in terms of incidental outcomes
Table XVII Reef Crest Foraging: Profitability and Overall Returns
Prey type Gross kcal Net kcal Encounter rate/min E/h
Turbo setosus N=2 57.5 SD=19.8 52.8 SD=19.5 0.2 SD=0.1 11.7 SD=3.2
E/T (kcal/min)
0.2
SD=0.1
7
A conservation ethic is synonymous with the philosophical under-
pinnings of Deep Ecology, whereby all living things are considered to
possess an inherent value (Sessions, 1998). A user-oriented
conservation strategy, in contrast, is rooted in anthropocentric ethical
norms (Zimmerman, 1998).
192 Hum Ecol (2007) 35:179194
stemming from social inequalities (Brower, 1983). For
Kiribati, however, with its more egalitarian social structure,
the major challenge stems from the erosion of customary
marine tenure, which together with high human population
growth, urban crowding, more efficient extractive technol-
ogies, and expanding market opportunities, has undermined
recent attempts to help conserve resources (Thomas, 2001a;
2001b).
Acknowledgment This paper draws on doctoral fieldwork in
Kiribati between 1993 and 1998. My thanks go to Douglas Bird for
sharing his data and thoughts on shellfish gathering from the
perspective of human behavioral ecology, Margo Wilson and Martin
Daly for suggestions to improve this manuscript, and the many I-
Kiribati for their patience and hospitality. The manuscript also
benefited from comments by three anonymous reviewers. The
research was supported by Sigma Xi (Grant-in-Aid of Research
# 2719 and # 8326), Conchologists of America, Inc., The Hawaiian
Malacological Society, and the University of Hawaii Arts & Sciences
Advisory Council.
References
Abbott, R. R., and Garcia, J. (eds.) (1995). Management Plan for
Tarawa Lagoon, Republic of Kiribati: Volume III. Management
Plan, BioSystems Analysis, Santa Cruz, California.
Adams, T., Dalzell, P., and Ledua, E. (1999). Ocean resources. In
Rapaport, M. (ed.), The Pacific Islands: Environment & Society,
Bess, Honolulu, pp. 366381.
Alcazar, S. N., and Solis, E. P. (1986). Spawning, Larval Development
and Growth of Tridacna maxima (Rding) (Bivalvia: Tridacni-
dae). Silliman Journal 33: 5673.
Alvard, M. S. (1993). Testing the Ecologically Noble Savage
Hypothesis: Interspecific Prey Choice among Piro Hunters of
Amazonian Peru. Human Ecology 21: 355387.
Alvard, M. S. (1994). Conservation by Native Peoples: Prey Choice in
a Depleted Habitat. Human Nature 5: 127154.
Alvard, M. S. (1995). Intraspecific Prey Choice by Amazonian
Hunters. Current Anthropology 36: 789818.
Anderson, A. (1979). Prehistoric exploitation of marine resources at
Black Rocks Point, Palliser Bay. In Leach, B. F., and Leach, H.
M. (eds.), Prehistoric Man in Palliser Bay, National Museum of
New Zealand Bulletin, No. 21, Wellington, pp. 4965.
Aswani, S. (1998). The Use of Optimal Foraging Theory to Assess the
Fishing Strategies of Pacific Island Artisanal Fishers: A Meth-
odological Review. Traditional Marine Resource Management
and Knowledge 9: 1926.
Barlow, K. R., and Metcalfe, D. (1996). Plant Utility Indices: Two
Great Basin Examples. Journal of Archaeological Science 23:
351371.
Beckerman, S. (1983). Carpe diem: an optimal foraging approach to
Bari fishing and hunting. In Hames, R., and Vickers, W. (eds.),
Adaptive Strategies of Native Amazonians, Academic, New York,
pp. 269299.
Begossi, A. (1995). Fishing Spots and Sea Tenure: Incipient Forms of
Local Management in Atlantic Forest Coastal Communities.
Human Ecology 23: 387408.
Bn, C., and Tewfik, A. (2001). Fishing Effort Allocation and
Fishermens Decision-making Process in a Multi-species Small-
scale Fishery Analysis of the Conch and Lobster in Turks and
Caicos Islands. Human Ecology 29: 157186.
Bettinger, R. L. (1991). Hunter-Gatherers: Archaeological and
Evolutionary Theory, Plenum, New York.
Bettinger, R. L., Malhi, R., and McCarthy, H. (1997). Central Place
Models of Acorn and Mussel Processing. Journal of Archaeo-
logical Science 24: 887899.
Bird, D. W. (1996). Intertidal Foraging Strategies among the Meriam
of the Torres Strait Islands, Australia: An Evolutionary Ecolog-
ical Approach to the Ethnoarchaeology of Tropical Marine
Subsistence, Doctoral dissertation, University of California,
Davis.
Bird, D. W. (1997). Behavioral ecology and the archaeological
consequences of central place foraging among the Meriam. In
Barton, C. M., and Clark, G. A. (eds.), Rediscovering Darwin:
Evolutionary Theory and Archaeological Explanation, Archaeo-
logical Papers of the American Anthropological Association No.
7, Arlington, pp. 291306.
Bird, D. W., and Bliege Bird, R. (1997). Contemporary Shellfish
Gathering among the Meriam of the Torres Strait Islands,
Australia: Testing Predictions of a Central Place Foraging Model.
Journal of Archaeological Science 24: 3963.
Bird, D. W., and Bliege Bird, R. (2000). The Ethnoarchaeology of
Juvenile Foragers: Shellfishing Strategies among Meriam Chil-
dren. Journal of Anthropological Archaeology 19: 461476.
Brower, K. (1983). A Song for Satawal, Harper Row, New York.
Catterall, C. P., and Poiner, I. R. (1983). Age and Sex-dependent
Patterns of Aggregation in the Tropical Gastropod Strombus
luhuanus. Marine Biology 77: 171182.
Charnov, E. L., and Orians, G. H. (1973). Optimal Foraging: Some
Theoretical Explorations. Manuscript on file, Department of
Biology, University of Utah, Salt Lake City.
De Boer, W. F., Blijdenstein, A.-F., and Longamane, F. (2002). Prey
Choice and Habitat Use of People Exploiting Intertidal Resour-
ces. Environmental Conservation 29: 238252.
Durnin, J. V. G. A., and Passmore, R. (1967). Energy, Work and
Leisure, Heinemann, London.
Dwyer, P. D. (1994). Modern Conservation and Indigenous Peoples:
In Search of Wisdom. Pacific Conservation Biology 1: 9197.
Emlen, J. M. (1966). The Role of Time and Energy in Food
Preference. American Naturalist 100: 611617.
FAO (1998). Fishery Country Profile: Kiribati, FAO, Rome.
FAO/WHO/UNU (1985). Energy and Protein Requirements, Techni-
cal Report Series No. 724, World Health Organization, Geneva.
Friedlaender, J. S., and Rhoads, J. G. (1987). Longitudinal anthropo-
metric changes in adults and adolescents. In Friedlaender, J. S.
(ed.), The Solomon Islands Project: A Long-term Study of Health,
Human Biology, and Culture Change, Clarendon, Oxford, pp.
283306.
Healey, C. (1993). The significance and application of TEK. In
Williams, N. M., and Baines, G., Traditional Ecological
Knowledge: Wisdom for Sustainable Development, Centre for
Resource and Environmental Studies, Australian National Uni-
versity, Canberra, pp. 2126.
Henry, C. J., and Rees, D. G. (1991). New predictive equations for the
estimation of basal metabolic rate in tropical peoples. European
Journal of Clinical Nutrition 45: 177185.
Jones, K. T., and Madsen, D. B. (1989). Calculating the Cost of
Resource Transportation: A Great Basin Example. Current
Anthropology 30: 529534.
Kaplan, H., and Hill, K. (1992). The evolutionary ecology of food
acquisition. In Smith, E. A., and Winterhalder, B. (eds.),
Evolutionary Ecology and Human Behavior, Aldine de Gruyter,
New York, pp. 167201.
Krebs, J. R., and McCleery, R. H. (1984). Optimization in behavioural
ecology. In Krebs, J. R., and Davies N. B. (eds.), Behavioural
Ecology: An Evolutionary Approach, 2nd ed., Blackwell,
London, pp. 91121.
Hum Ecol (2007) 35:179194 193
Leung, W.-T., Butrum, R. R., and Chang, F. H. (1972). Proximate
composition, mineral and vitamin contents of East Asian foods.
In Food Composition Table for Use in East Asia, U. S.
Department of Health, Education, and Welfare, Bethesda, pp.
1187.
MacArthur, R. H., and Pianka, E. A. (1966). On Optimal Use of a
Patchy Environment. American Naturalist 100: 603609.
Marsh, J. A., Jr. (1987). Reef processes: Energy and materials flux. In
Devaney, D. M., Reese, E. S., Burch, B. L., and Helfrich, P.
(eds.), The Natural History of Enewetak Atoll: Volume I. The
Ecosystem: Environments, Biotas, and Processes, U. S. Dept. of
Energy, Office of Energy Research, Office of Health and
Environmental Research, Ecological Research Division, Oak
Ridge, pp. 159179.
Maynard Smith, J. (1978). Optimization Theory in Evolution. Annual
Review of Ecology and Systematics 9: 3156.
McNab, B. K. (1963). Bioenergetics and the Determination of Home
Range Size. American Naturalist 97: 133140.
McNeely, J. A., Houston, A. I., and Weisser, W. W. (1995). Human
influences on biodiversity. In Heywood, V. H., and Watson, R. T.,
Global Biodiversity Assessment, Cambridge University Press,
Cambridge, pp. 715821.
Metcalfe, D., and Barlow, K. R. (1992). A Model for Exploring the
Optimal Trade-off Between Field Processing and Transport.
American Anthropologist 94: 340356.
Mitchell, D., Nash, J., Ogan, E., Ross, H., Bayliss-Smith, T., Keesing
R., and Friedlaender, J. S. (1987). Profiles of the survey sample.
In Friedlaender, J. S. (ed.), The Solomon Islands Project: A Long-
term Study of Health, Human Biology, and Culture Change,
Clarendon, Oxford, pp. 2864.
Murai, M. Pen, F., and Miller, C. D. (1958). Some Tropical South
Pacific Foods: Description, History, Use, Composition, and
Nutritive Value, University of Hawaii Press, Honolulu.
Norgan, N. G., Ferro-Luzzi, A., and Durnin, J. V. G. A. (1974). The
Energy and Nutrient Intake and the Energy Expenditure of 204
New Guinean Adults. Philosophical Transactions of the Royal
Society of London, Series B, 268: 309348.
Nydon, J., and Thomas, R. B. (1989). Methodological procedures for
analyzing energy expenditure. In Pelto, G. H., Pelto, P. J., and
Messer, E. (eds.), Research Methods in Nutritional Anthropology,
United Nations University, Tokyo, pp. 5781.
Pargeter, K, Taylor, R., King, H., and Zimmet, P. (1984), Kiribati: A
Dietary Study, Noumea, South Pacific Commission.
Paulay, G. (2001). Benthic Ecology and Biota of Tarawa Lagoon:
Influence of Equatorial Upwelling, Circulation, and Human
Harvest. Atoll research Bulletin 487: 141.
Pernetta, J. C., and Hill, L. (1984). Traditional Use and Conservation
of Resources in the Pacific Basin. Ambio 13: 359364.
Pyke, G. H., Pulliam, H. R., and Charnov, E. L. (1977). Optimal
Foraging: A Selective Review of Theory and Tests. Quarterly
Review of Biology 52: 137154.
Redford, K. H. (1991). The Ecologically Noble Savage. Orion 9: 2429.
Ruddle, K., Hviding, E., and Johannes, R. E. (1992). Marine
Resources Management in the Context of Customary Tenure.
Marine Resource Economics 7: 249273.
Ruttan, L. M., and Borgerhoff Mulder, M. (1999). Are East African
Pastoralists Truly Conservationists? Current Anthropology 40:
621652.
Salvat, B. (1972). La faune benthique de lAtoll de Reao (Tuamotu,
Polynsie). Cahiers du Pacifique 16: 31110.
Sessions, G. (1998). Introduction. In Zimmerman, M. E. (ed.),
Environmental Philosophy: From Animal Rights to Radical
Ecology, 2nd ed., Prentice Hall, Upper Saddle River, N. J., pp.
165182.
Sidwell, V. D., Foncannon, P. R., Moore, N. S., and Bonnet, J. C.
(1974). Composition of the Edible Portion of Raw (Fresh or
Frozen) Crustaceans, Finfish, and Mollusks. I. Protein, Fat,
Moisture, Ash, Carbohydrate, Energy Value, and Cholesterol.
Marine Fisheries Review 36(3): 2135.
Smith, E. A. (1983). Anthropological Applications of Optimal
Foraging Theory: A Critical Review. Current Anthropology 24:
625651.
Smith, E. A. (1991). Inujjuamiut Foraging Strategies: Evolutionary
Ecology of an Arctic Hunting Economy, Aldine de Gruyter, New
York.
Smith, E. A. (1995). Comments on Intraspecific Prey Choice by
Amazonian Hunters by M. Alvard. Current Anthropology 36:
810811.
Smith, E. A., and Winterhalder, B. (1992). Natural selection and
decision making: Some fundamental principles. In Smith, E. A.,
and Winterhalder, B. (eds.), Evolutionary Ecology and Human
Behavior, Aldine de Gruyter, New York, pp. 2560.
Sosis, R. (2002). Patch Choice Decisions among Ifaluk Fishers.
American Anthropologist 104: 583598.
Stephens, D. W. (1985). How Important are Partial Preferences?
Animal Behaviour 33: 667689.
Stephens, D. W., and Krebs, J. R. (1996). Foraging Theory, Princeton
University Press, Princeton.
Sutherland, W. J. (1982). Do Oystercatchers Select the most Profitable
Cockles? Animal Behaviour 30: 857861.
Tamaru, C. S., Ako, H., and Lee, C.-S. (1992). Fatty Acid and Amino
Acid Profiles of Spawned Eggs of Striped Mullet, Mugil
cephalus L. Aquaculture 105: 8394.
Thomas, F. R. (1999). Optimal Foraging and Conservation: The
Anthropology of Mollusk Gathering Strategies in the Gilbert
Islands Group, Kiribati. Doctoral dissertation, University of
Hawaii, Honolulu. University Microfilms International, Ann
Arbor.
Thomas, F. R. (2001a). Mollusk Habitats and Fisheries in Kiribati: An
Assessment from the Gilbert Islands. Pacific Science 55: 7797.
Thomas, F. R. (2001b). Remodeling Marine Tenure on the Atolls: A
Case Study from Western Kiribati, Micronesia. Human Ecology
29: 399423.
Thomas, F. R. (2002). An Evaluation of Central-place Foraging
among Mollusk Gatherers in Western Kiribati, Micronesia:
Linking Behavioral Ecology with Ethnoarchaeology. World
Archaeology 34: 182208.
Ulijaszek, S. J. (1995). Human Energetics in Biological Anthropology,
Cambridge University Press, Cambridge.
Villiers, L., and Sire, J-Y. (1985). Growth and Determination of
Individual Age of Turbo setosus (Prosobranchia Turbinidae), Hao
Atoll (Tuamotus, French Polynesia). Proceedings of the Fifth
International Coral Reef Congress 5: 165170.
Wiens, H. J. (1962). Atoll Environment and Ecology, Yale University
Press, New Haven.
Yamaguchi, M., Yeeting, B., Toyama, K., Tekinaiti, T., and Beiatau, T.
(1993). Report on a Preliminary Study of Molluscan Resources in
Tarawa Lagoon, with Special Reference to te bun (Anadara
maculosa [sic]), Department of Marine Sciences, University of
the Ryukyus, Okinawa.
Zimmerman, M. E. (1998). General introduction. In Zimmerman, M.
E. (ed.), Environmental Philosophy: From Animal Rights to
Radical Ecology, 2nd ed., Prentice Hall, Upper Saddle River, N.
J., pp. 16.
194 Hum Ecol (2007) 35:179194