ANNUAL

REVIEWS Further
Quick links to online content

Ann. Rev. Entomol. 1989. 34:191-210

Copyright © 1989 by Annual Reviews Inc. All rights reserved

FORAGING STRATEGIES OF ANTS

by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org

James F. A. Traniello

Department of Biology, Boston University, Boston, Massachusetts 02215

OVERVIEW: ANT FORAGING STRATEGY AS AN

INDIVIDUAL AND SOCIAL PROCESS

Ant foraging is a collective process composed of the activities of individuals

as well as behaviorally integrated groups. Therefore, a great challenge in the
socioecology of foraging is to explain how the behavior of such a potentially
large and complex system as an ant colony emerges as a function of the
properties of its individual components. The task of studying foraging in ants
in simplified by the fact that foragers, owing to their sterility, may do little
more than forage during their tenure as food harvesters. However, eusociality
penetrates virtually every facet of foraging strategy and adds additional
complicating social dimensions to its analysis.
Individual and social processes of foraging can be categorized into com­
ponents to facilitate the analysis of foraging at the level of the single worker,
the colony, or the integration of both (Table 1). A generalized ethogram of
forager behavior can be described to illustrate these processes. A forager
leaves the nest entrance in either a random or a consistent direction. A travel
phase ensues, during which the worker maintains a constant compass bearing
and moves directly away from the nest. At some point during the travel phase
the forager shows a high frequency of turning, marking the beginning of
search. During search, food resources are encountered and selected based
upon a forager's physical caste, age, and prior experience, the trip distance,
thermal stress, resource quality, and the colony's current nutritional status.
Depending on the ant species and the size, density, or quality of the food, the
forager may communicate information to nestmates about its location and
recruit additional foragers.
Summing the above-described individual activities across all foragers
yields the most basic aspect of cooperative foraging. Changing nutritional
demands associated with the maturation of reproductives may shift a colony's

191
0066-4170/89/0101-0191$02.00
 192 TRANIELLO

Table 1 Individual and social c?mponents of ant foraging systems, with representative refer­
ences

Individual Social

I. Trip direction, distance, and duration 1 . Summation of individual efforts (41,

(92,1 12,1 1 8, 1 1 9) 1 1 9)
2. Search pattern and duration (91, 1 1 2 , 2. Temporal and physical caste; colony
1 1 8, 1 1 9) demographic distribution ( 1 2,24, 33,
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.

59, 75 , 78)
3. Loading capacity and retrieval (3, 34, 3. Tempo (60, 75)
40,63,73, 89)
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org

4. Resource assessment and selection (6, 4. Communication and cooperative re­

n.�,M,m.�,�.�.�.�, trieval (I, 34, 50, 53, 1 09, 1 1 0)
1 00, 1 1 6, 1 1 7, 1 1 9)
5 . Induction of cooperative foraging (I, 5. Resource assessment and colony regu­
19, 109) lation of foraging (96, 1 00, 1 0 1 , 1 04-
1 06) (see also first column, component
4)

foraging actIvlty among different food types. Communication mediates

cooperation among foragers during search and resource retrieval and serves as
a control mechanism of colony-level foraging responses. As a result it pro­
vides regulatory feedback during the daily and seasonal organization of
foraging.
In this review I treat foraging strategy as a system divisible into elements
composed of individual and collective action, recognizing that the distinction
between the two may often be blurred because of the high degree of social
integration of colony foraging activity. My goal is to provide a synthesis of
selected current literature to supplement available reviews (9-11, 14, 43, 57,
75, 76, 98, 120).

FORAGING SYSTEM EVOLUTION: A COMPONENT

ANALYSIS APPROACH

the traits that are used to categorize ant foraging are highly variable in their
distribution among the more primitive (e.g. ponerine) or advanced (e.g.
forrnicine) subfamilies of ants. For example, "diffuse foraging" (75), in
which foragers leave the colony singly and retrieve food solitarily, occurs in
the Ponerinae and Myrrneciinae. The forrnicines Cataglyphis spp. and
Ocymyrmex spp. also collect randomly dispersed food largely through in­
dividual effort (l18, 119), but the ponerines Pachycondyla laevigata and
Leptogenys spp. have chemically mediated cooperative foraging (54, 68).
Ecological influences on foraging systems, therefore, seem to override
phylogenetic tendencies. Also, any one foraging mode must not be mistaken
 ANT FORAGING STRATEGIES 193

for a species-typical characteristic when in fact it may be a behaviorally

flexible component of total foraging strategy. The use of such a foraging
categorization to indicate the outcome of higher-level ecological processes
may thus yield conclusions that are not entirely valid. For example, the gross
categorization of ant species as being "individual," "recruit," and/or "group"
foragers (4, 5) may be "too weak and artificial to be useful" (52) when used to
define foraging systems in studies of community organization. Given the wide
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.

variety of feeding habits and levels of organization and complexity, I suggest

that foraging systems be analyzed as a series of components that may each
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org

have individual and social influences and that have all evolved in response to
resource distribution patterns, competition, and predation. The designation of
systems or species as "individual foraging" should be dropped because the
behavior of a forager is probably never completely independent of the activi­
ties of other foragers and the state of the colony as a whole. Therefore, the
idea of individual foraging is erroneous or at least misleading, even in species
lacking recruitment communication. Some components of foraging behavior,
such as search or retrieval, may largely be individual activities. Search and
retrieval tactics will depend on the size of resources, their temporal and spatial
distribution patterns, and their resistance to retrieval, as well as on the loading
capacities of foragers. Therefore, retrieval may be an individual or group
process when viewed as a component of foraging, but this one feature should
not be used to categorize the entire system. In summary, any single
categorization of ant foraging systems is inadequate in that it does not
consider the components of foraging behavior and their ecological influences,
which are necessary to fully understand the total expression of a colony's
foraging strategy.

Caste Evolution and Foraging Strategy

Because foragers are sterile they do not have conflicting demands on their
time and energy budgets that involve trade-offs between searching for food
and searching for mates. As a result, they are often viewed as the products of
selection for traits that maximize energy return to the colony. Several social
traits have consequences that affect a colony's economy; among these are
temporal polyethism and senescence, caste polymorphism, and foraging tem­
po (60, 75, 78). Selection at the colony level can not only affect the be­
havioral repertories of foragers to enhance their food harvesting ability, but
can also shape patterns of age demography and mortality. In Pogonomyrmex
owyheei. the life expectancy of a forager averages 14 days under natural
conditions; survivorship is similar in the laboratory under conditions of
starvation (78). However, in Cataglyphis bicolor, foragers have a life ex­
pectancy of 6.1 days in the field but live for months or longer in the laboratory
(93). Colony fitness may be correlated with the morphology and size frequen-
194 TRANIELLO

cy distribution of foragers. Their structural morphology per se may reflect

feeding habit. Although it is clear in the case of some specialist predators such
as Amblyopone, Odontomaehus, and Strumigenys species that mandibular
shape and head structure reflect feeding habits, the relationship between
morphology and foraging ecology is often unclear. Recent studies using
digitization analysis to detect the ecological correlates of variation in head
morphology have shown that head shape, as differentiated from size, is
related to diet (B. Cole, personal communication). Other studies have
ascribed a role to vertebrate pred �tion as a selective agent of form (56).

Caste, Competition, and Community Organization

Among monomorphic species, body size may shift owing to interspecific
competition and may facilitate coexistence. It is assumed that there is a
correlation between forager size and food-item size, so that morphological
divergence produces resource partitioning. Body size of C. bieolor foragers is
considerably smaller in populations in Greece than in Tunisia, apparently
owing to the sympatry of the smaller-bodied Cataglyphis albieans in the latter
environment (119). In Greece, where C. albieans is absent, the body size of
C. bicolor foragers is intermediate in size between that of C. albicans and C.
bicolor in Tunisia, which suggests that a morphological shift has occurred
because of local competition. Several studies have indicated a correlation
between body size and food-item size for desert seed-eating ant communities
and communities of generalist ant species (16, 17, 21, 38, 75, 110, 1 18).
However, given the limitations of the use of body size as an indicator of food
choice and niche (46, 47, 122) and the fact that the food recruitment systems
of ants make them less constrained by their morphology than other animals
(14, 20, 1 10), size-based partitioning of resources among sympatric species
may be the exception rather than the rule in ants (2 1).
The species diversity of granivorous ants of the southwest desert in North
America is related to productivity, and coexistence is based upon body size
differences or foraging strategies that are correlated with seed size selection or
seed density distributions (2 1, 22). In the Australian arid zone, species
density is similar to that of North American deserts, but there is less separa­
tion by size among sympatric species (72). Also, differences in productivity
among sites do not appear to have a strong influence on associated harvester
ant communities in Australia as in North America, seemingly because popula­
tion density is not limited by food availability (7, 8). Vegetation structure,
and perhaps productivity, appears to be related to community organization (2,
7, 8, 25). Worker size/seed size correlates can be found among some species,
but other factors such as seed morphology and biochemistry affect seed choice
as well (25). In the Negev Desert one of three sympatric Messor species
shows a small but significant correlation between forager size and seed size
 ANT FORAGING STRATEGIES 195

harvested (S. Rissing, personal communication). In North-Temperate open

field ants and other communities of ants with similar generalist feeding habits,
prey size is correlated with body size but mechanisms of cooperative foraging
and interference also seem to be important determinants of prey selection and
community-level interactions (1, 22, 29, 31, 49, 53, 65, 109, 110).

Caste Polymorphism and Foraging Ecology

Although it occurs in a small fraction of ant species, the role of caste
polymorphism in foraging has received much attention. Has natural selection
adaptively shaped colony demography? Do the proportions of workers of
different size castes match the characteristics of food resources, and if so,
how closely (75)? Size matching, the pairing of forager size and food-item
size, has been suggested for a number of species (21, 24, 75, 80), but the
correlation explains little of the variance in food choice (83, 84). Competi­
tion, diet expansion, and polymorphism might be linked in Pogonomyrmex
spp. (75), but in the polymorphic Pogonomyrmex badius there is only a low,
insignificant correlation between polymorphism and diet (1. F. A. Traniello &
S. N. Beshers, unpublished). Colonies of Veromessor pergandei show in­
creased size variation of workers in areas of relaxed interspecific competition
(24). However, less than 4% of the variance in the size of seeds collected by
V.pergandei is explained by forager body size, and worker body size changes
during seasonal cycles, showing little correlation with seed size or the per­
formance of any task (83, 84). In this species, worker size may change
according to levels of resource availability, usable foraging time, and alate
production, rather than according to selection for size pairing (83).
Polymorphism in Edton burchelli is thought to enhance foraging efficiency
by lowering prey retrieval costs. Submajors, which comprise 3% of the
colony, are disproportionately active in prey transportation, representing 26%
of all prey carriers (33). Leg length allometry may increase foraging velocity.
Across physical castes, prey dry weight is significantly correlated with worker
dry weight; submajors carry the largest prey loads, although loaded submajors
do not transport prey more rapidly than other castes (33). The inverse
relationship between ant body size and respiratory rate (3, 63) and computer
simulations (80) support the notion of enhanced foraging efficiency in a
polymorphic series, but critical demonstrations are lacking.

FORAGER LOAD CAPACITY AND CASTE EFFICIENCY One of the most

striking features of ant biology that bears on foraging efficiency is the ability
of an individual forager to carry a resource load that is considerably heavier
than its own body weight. The loading ability of a forager, measured as
burden (APnt + Mload/Mant, where M is mass (3, 82), is 1.5-7.5 in Atta
cephalotes (89), 1.2-1.5 in E. burchelli (34), on average 2.2 in Pogono-
196 TRANIELLO

myrmex occidentalis (32), and 1.1-15:3 in Formica schaufussi O. F. A.

Traniello, unpublished.) Eciton burchelli submajors have disproportionately
long legs and mandibles, and can easily straddle and transport prey. While
this mode of rapid prey transport appears efficient, the burden capacity of lone
foragers is greater in species such as F. schaufussi that drag rather than lift
their prey. In this latter case prey transport speed decreases sharply with
increasing load weight, most likely because of frictional resistance (J. F. A.
Traniello, unpublished). In polymorphic species, large workers may carry
larger loads, but the cost of producing these workers and the energetic gain
from their input to a colony's nutritional economy are unclear. For example,
in Eciton spp. increasing forager weight increases load capacity and possibly
efficiency (12, 34). However, a standard calculation of burden reveals load
ratios are 1.2, 1.3, and 1. 5, respectively, for 2.5-,5-, and lO-mg workers;
therefore, larger workers have only slightly greater load capacities. Transport
costs decrease with increasing body size in ants in general, and in Eciton
hamatum the gross and net costs of transport decrease with body mass and
running speed (3). However, the cost for polymorphic foragers of transporting
average-size burdens must be estimated before transport efficiency can be
fully evaluated.

RESOURCE DISTRIBUTION, POLYMORPHISM, AND COOPERATIVE RE­

TRIEVAL A significant body size/prey size correlation does not necessarily

imply that competition has led to divergence in worker morphology. First,
morphology permits a worker to collect not only prey matched to its body
size, but also any smaller load as well. This conclusion is supported by the
fact that for some species variance in prey size increases as body size
increases (110). Second, the range of prey sizes available or actually taken
may be extremely large in comparison to the range of worker body sizes in
some sympatric ant species, so body size variation can at best adapt a colony
to collecting a limited range of available prey. Third, recruitment communica­
tion, which is common in many species, provides a flexible mechanism for
simulating a larger-bodied forager by organizing a cooperative retrieval group
(34, 53, 109). Given the unpredictable distribution patterns of the food
sources of many ant species, the flexibility of caste simulation through
cooperative retrieval appears to best adapt a colony to temporal, spatial, and
size variability; resource distribution patterns may vary on a relatively brief
ecological time scale, but worker size demography cannot be shifted within
the same time frame.
Cooperative prey retrieval occurs in many species of ants, but its evolution,
mechanics, and energetics are poorly understood. In Lasius neoniger. F.
schaufussi. and E. burchelli, transport group size is correlated with prey
weight. seemingly to maximize carriage efficiency (34. 109; J. F. A. Tra-
 ANT FORAGING STRATEGIES 197

niello, unpublished). In E. burchelli, groups are structured teams composed

of major workers plus one submajor; this group composition seems to
minimize rotational forces during prey movement (34). In other ants, howev­
er, groups often seem to lack structure beyond a forager number/prey weight
relationship and to lack coordination. In F. sc ha ufussi only some of the ants in
a recruited group are involved in load carriage; some workers ride atop the
prey while others pull in opposing directions. The loading capacity of coop­
erative retrieval groups ranges from 1. 7 to 8.9, which is lower than the
loading capacity of single workers. In this species, and in Novomessor spp. ,
cooperative retrieval may be organized to decrease interference competition
through resource defense rather than to move prey with greater energetic
efficiency (J. F. A. Traniello, unpublished).

FORAGING THEORY AND DIET SELECTION

It has been argued that ants are ideally suited to test predictions of theories of
feeding strategies because of their eusociality; natural selection has made
foragers efficient food harvesters that maximize a colony's energy intake
(75). Equally convincing is the argument that difficulties arise in the applica­
tion of optimization models to ants because the forager population of a colony
is often large and sometimes polymorphic, and worker foraging activity is
severely constrained by temperature (80). Although there have been many
demonstrations of food selection in ants (6, 23,30,55,64,70,79,81,86,
89, 92, 99, 100, 116, 117, 119), tests of theory have shown varying degrees
of correspondence between ant foraging behavior and the predictions of
models (23, 48, 94, 100). The seed-harvester ants Pogonomyrmex rugosus
and Pogonomyrmex barbatus increased diet specialization with increasing
distance of seed patches from the nest, but showed partial preferences (23).
Similar tests conducted with P. occidentalis and Pogonomyrmex californicus
showed no significant change or a nonsignificant trend in seed size choice as a
function of distance (48,100). However, distances at which seed patches were
offered were variable for the different species (3-12 m vs 3-7 m), and for P.
californicus the maximum distance was 85 cm from the nest. The basic
premise of these studies is that the net energy value of seeds of all sizes
decreases with increasing distance from the nest owing to the added cost of
travel and transport, so that energy maximization is achieved at the more
distant patches through the selection of large seeds. The assumption is that
energetic costs of search or travel and transport are significant relative to
energy gain. However, foraging costs for P. occidentalis workers are less
than 0.1% of the caloric content of an average collected seed (32). This
suggests that feeding selectivity may not change substantially with increasing
198 TRANIELLO

travel costs and that time cost and net energetic gain rate may be important
criteria in foraging strategy (32).
Ants are poikilothermic, and their foraging activity is therefore constrained
by temperature, water stress, and other physical factors that might affect the
energetic costs of foraging or the use of time. Several studies have demon­
strated a significant effect of ambient temperature on metabolic rate as
measured by oxygen consumption (26, 77). In F. schaufussi the elevated rate
of oxygen consumption appears to be an important component of foraging
cost that influences prey selection (114). The role of travel distance and its
importance relative to temperature as a foraging cost has also been examined
in F. schaufussi. Prey of two different sizes were offered at distances of I and
5 m from the nest at low and high temperatures. At both 1 and 5 m, increase in
temperature produced a decrease in selectivity; increasing travel costs at either
low or high temperature, however, did not affect selectivity (R. V. Bowen &
J. F. A. Traniello, unpublished). The magnitudes of temperature-induced
costs and travel costs seem therefore to be very different; temperature pro­
duces the more important constraint, if in fact temperature represents a
foraging cost rather than a thermal risk factor.
Laboratory studies of patch use by Solenopsis geminata have shown that
colonies recruit more foragers to sucrose patches that are closer to the nest,
larger, or of higher concentration than to more distant, lower-quality, or
smaller patches (100). Patterns of recruitment and patch use in S. geminata
support models of energy maximization rather than time minimization. Sim­
ilar support for the hypothesis of energy maximization is provided by P.
rugosus and P. barbatus, which recruit more foragers to seed patches close to
the colony (100).
Leafcutting ants select tree species based on the presence or absence of
secondary plant compounds, water content, and leaf density (6, 55, 86,116,
117). Colonies are more selective of species the more distant they are from the
nest, although at these foraging loci they sometimes do not harvest the leaves
of neighboring palatable species (86). Foraging by leafcutting ants follows the
prediction of optimal diet theory, which states that colonies should show
greater diet diversity and lower resource constancy when average resource
quality is low; colonies indeed shift their leafcutting among a greater number
of tree species when the average palatibility of resource trees is low, and
specialize in richer-resource environments (86, 95). Also,70% of observed
colonies cut relatively close to the nest, and foragers cut leaves from higher­
quality tree species with increasing distance from the nest (86).
Risk as a factor influencing foraging decisions has been examined through
studies of both simulated and natural predation (37,66,121) and competition
(49, 113), which result in a decrease or cessation of foraging or a change in
foraging direction. The influence of competitors on the spatial patterning of
 ANT FORAGING STRATEGIES 199

foragers and resource use has been described in Pogonomyrmex spp. (49) and
in L. neoniger (113). In these species confrontation between foragers from
adjacent colonies leads to altered trail system directionality and search pat­
tern, respectively, without a reduction in overall foraging activity. In Lasius
pallitarsus, resource use (patch choice) is affected by the presence of Formica
subnuda workers; L. pallitarsus workers shift foraging activity to a safe,
competitor-free patch, even if it is of lower quality (74).
Theories of foraging strategy provide an important organizational
framework with an emphasis on energetics that is valuable for the study of the
foraging economy of ants. But optimization models should be cautiously
applied because the economics of foraging cannot be studied in isolation from
other aspects of ecology, energetics, social organization, and the intrinsic
behavioral limitations of ants. For example, some models (97) assume forager
knowledge of the resource base as a requirement for optimal feeding strategy,
yet the relatively short life span of a forager may limit an individual ant's
experience. Moreover, if resources are unpredictably distributed, it is difficult
to see how such information would modify future individual or colony
decisions. The spatial pattern of foraging may also limit the acquisition of
information about resource distribution. Low encounter rates and foraging
success may be additional complications. Little is known of the memory
capability of ants in rcgard to food choice; a fcw studies show they have the
ability to store information and use it in foraging decisions (81, 87, 114; R.
Johnson, personal communication). Should foragers have knowledge of re­
source availability, how are the preferences of individuals summed to yield a
colony-wide pattern of choice? The role of these factors in the implementation
of foraging strategy is poorly understood.
The social organization of foraging may influence spatial patterns of search
and therefore resource assessment. The trunk trail systems of Atta spp., for
example, originate from pheromone deposition and guide foragers to a tree to
be attacked; ultimately, a path between the nest and the tree is cleared of
obstruction. While this path enhances leaf transport efficiency, it limits ants to
searching for new host trees only near the trunk route's end and sides (86, 94).
Similarly, the use of paths cleared of vegetation by P. occidentalis foragers
enhances gain rate but probably canalizes search (32).

SEARCH PATTERN

Search pattern can be analyzed at two levels. First, search concerns the use of
foraging space by colonies within populations, and intra- and interspecific
territorial interactions may contribute to the partitioning of space between
colonies (42, 49, 51, 62, 90, 113, 115). Therefore, search pattern involves
colony-level regulation of the spatial allocation of foragers. Second, each trip
200 TRANIELLO

made by a forager has its own ecologically influenced spatial and temporal
organization (41, 91, 112, 118, 119). There is also a relationship between
individual and colony-wide search patterns, since a forager may influence the
search patterns of nestmates through communication. At both levels, resource
distribution in space and time and competition should be major influences on
the organization of search.

Territoriality and Search Pattern

The significance of the spacing patterns of ant colonies and the division of
foraging area has chiefly been approached by measuring dispersion and/or by
studying territorial interactions (51, 62). Cost/benefit explanations suggest an
interplay of the economics of foraging and defense and allow predictions to be
made about how colonies should allocate workers to related tasks in time and
space (51, 58). Dispersion patterns may change seasonally and appear to be
nonrandom in most species for which data are available (42, 45, 49, 62, 90,
115).
Trail systems provide a mechanism by which the search patterns of foragers
are directed away from neighboring colonies (42, 49, 90, 113). They also
reduce search or travel time by directly channeling foragers to a persistent
resource (49, 94, 95). In some cases more than one function may be served.
Colonies of P. rugosus and P. barbatus feed at seed patches, and foragers
travel along spatially separated trunk trails; the average nearest-neighbor
distance between nests is 17-18 m (49). In contrast, Pogonomyrmex marico­
pa takes evenly distributed seeds, and foragers seem to depart randomly from
the nest and collect seeds individually; colonies of this species are on average
46 m apart, apparently owing to the absence of trail systems (49). In C.
bicolor, the radius of an area in which 90% of all search activity occurs
approximates the average internest distance (119). In Pheidole militicida,
trunk routes branch into smaller trails, and the persistence of use of these
components of the trail system varies, with the trunk being most stable (52).
Trunk routes in P. militicida channel foragers to areas where seeds are
available, and the routes are abandoned when the resources are depleted. The
biochemical ecology of trail system formation has been studied in regard to
level of specificity and forager orientation (51, 52, 108, 113). Trail pher­
omones may reduce search time by serving as initial guides to areas of high
seed density and may also perhaps decrease homing time, although visual
cues are also important in orientation (49, 113). In L. neoniger the colony­
specific persistent components of trail pheromone appear to direct forager
search paths away from neighboring colonies, which may be as close as 5 cm
(l08, 113, 115). The highly directional patterns of foraging become random
following confrontation with foragers from an alien nest, which suggests that
the trail systems of this species minimize aggressive confrontation with
competitors (113).
 ANT FORAGING STRATEGIES 201

Geometry of Search at the Colony Level

In the desert harvester ant V. pergandei the direction of foraging columns is
rotated in a clockwise or counterclockwise manner between consecutive
foraging periods, and the size of a foraging column is adjusted to seed
production and patch distance (85). As food density decreases, columns
increase in length and rotate more frequently. Column rotation seems to
represent a system of sectorial search with a radial shift following resource
depletion. Consecutive raids of the army ant E. burchelli are systematically
rotated an average of 1230 during the statary phase (35). Raiding also appears
to be systematic in the nomadic phase, again seemingly for avoidance of
redundant search. The geometry of search appears to be determined solely by
prior raid direction, perhaps with modifications to avoid contact of foragers
with other E. burchelli colonies (35).
Such systematic search during the statary and nomadic phases does not,
however, appear to be the rule among army ants. In Neivamyrmex nigrescens
the directions and distances of emigrations vary unpredictably during the
nomadic phase (71). The striking differences in the search and raiding pat­
terns of E. burchelli and N. nigrescens may be related to the distribution and
abundance of prey or to nest sites and climatic factors (71). However, the
density of prey species of ants in the southwest desert or grassland environ­
ment of N. nigrescens is approximately 3000 colonies/ha (71), and the density
of ant species actually or potentially taken by E. burchelli is on average 6600
± 2200 colonies/ha (range 4000-9000 colonies/ha) (61). The highly orga­
nized raid pattern of E. burchelli may therefore reflect selection for efficient
prey harvesting given the greater abundance of its prey, but the role of prey
distribution is not clear. Limited study of the nomadic obligate termite
predator Pachycondyla (=Termitopone) laevigata has shown no predictable
raiding pattern (30).

Search Patterns of Individual Foragers

Forager decisions may include where to search, how long to search at a given
site, and whether or not to return to a site where search was previously
conducted. Some species, such as P achycondyla ( N eoponera) apicalis,
=

show strong route fidelity, each worker restricting its foraging activity to a
given site (36). In C. bicolor search effort is concentrated in sectors of
approximately 50° and 30-m radius, and average search path length differs
among populations according to food availability (119). The search pattern of
a colony contains random components as evidenced in the exponential dis­
tribution of search times of foragers, increasing average distance of search in
proportion to the square root of time, and selection of initial foraging direc­
tion. Thus the colony-level search pattern approximates a random diffusion
process, which may be an efficient solution to harvesting food that is un­
predictably distributed (41). In F. schaufussi, search pattern is related to food
202 TRANIELLO

distribution; collecting a single crop load of carbohydrate food produces a

more spatially restricted and accurate search during a return trip than collect­
ing a single load of insect prey (J. F. A. Traniello & A. J. Kozol, un­
published).

SEARCH DURAnON AND PERSISTENCE Search by C. hieolor foragers

shows distinctive temporal patterns that are correlated with food availability.
Colonies in Greece collect less prey biomass than colonies in Tunisia, and
individuals in Greece make roughly double the number of trips, although trip
duration is shorter (119). Trip duration is also dependent upon the type of
food collected; it is longer for ants collecting plant secretions (39).
The tendency of a forager to return to and search at a site where it has found
food has been studicd in C. hieolor (91) and F. sehaufussi (111, 112). In both
species foragers differ in their tendency to restrict their search to the site of a
prior find. Persistent foragers show intensive local search, whereas nonpersis­
tent foragers often completely bypass a prior reward site. In C. hieolor these
individually different search strategies appear to categorize foragers through­
out their lives and reflect foraging specializations. Search pattern in F.
sehaufussi appears to be adjusted in response to food type, and search
persistence seems to be correlated with the spatial and temporal persistence of
the insect prey and homopteran excretions that compose this species' diet
(12). Consecutive rewards of carbohydrate food tend to produce a shift from
nonpersistent to persistent searching and increase search duration (112). The
degree of spatial accuracy of search is greater in response to feeding on
carbohydrate than on insect prey, but it does not improve with successive
multiple rewards of either type of food, which indicates that spatial patterning
of search and information acquired to control search are fixed resource­
dependent parameters. However, search duration may be conditioned and
may at least have a local effect on persistence. In C. hieolor. search persis­
tence increases with the distance of the food to the nest (91), and in V.
pergande; the frequency of seed sampling in a patch increases with the
distance of the patch from the nest (84). Increasing search time in V. per­
gandei results in the sampling of a larger number of seeds in a patch.

SOCIAL REGULATION OF FORAGING

Age Polyethism and Development of Foraging Behavior

Workers of many ant species sequentially change tasks as they mature,

beginning with brood care following worker eclosion and culminating with
foraging (59, 75). Several factors affect the onset of foraging behavior. The
completion of physiological processes related to cuticular sclerotization and
the acquisition of sensory receptor competence may be required before forag-
 ANT FORAGING STRATEGIES 203

ing can begin. A callow phase during which tanning takes place may be
necessary to provide time for the completion of physiological maturation;
these processes occur during pupal development in some species (107).
Feeding habits could affect the pattern of age polyethism if brood care is
directly associated with foraging. This may be true for species such as
Amblyopone paUipes that directly provision larvae with prey (107). In the
temporal polyethism of most species, tasks with high associated risks, such as
foraging, are scheduled as the ultimate worker contribution to a colony's
economy (75). Ergonomically, loss is minimized if forager mortality is high
because foragers hold fewer energy reserves in the form of lipid stores than
younger workers (67).
The percentage of foragers in a colony varies across species (67, 78). The
theory of adaptive demography predicts that the proportion of workers allo­
cated to foraging should differ among populations in different ecological
circumstances if worker age determines the transition to foraging performance
(75). Recent field-based empirical tests suggest that age caste ratios are highly
variable, and the proportion of older workers in a colony shows no clear
correlation with ecology. Censuses of whole, field-collected Pheidole dentata
colonies from two habitats showing significant differences in food resource
biomass, ant species diversity, predation, and competitive environment re­
vealed that older workers, which are typically foragers, make up a varying
proportion of the total worker force (P. Calabi & J. F. A. Traniello, un­
published). Although temporal caste ratios vary, colonies with very different
demographic profiles have very similar labor profiles. This suggests that
changing foraging requirements are met by rapid changes in the task perfor­
mance patterns of individual workers in response to colony need rather than
by changes in age demography. This intercolony consistency in the proportion
of total labor involved in foraging suggests a nexible switching of the
behavior of individual workers in response to immediate foraging require­
ments. A worker's shift from another task to foraging is therefore related not
only to age but also to the perception of social stimuli carrying information
about the current nutritional status of a nest and/or the activities of other
colony members. In Novomessor albisetosus, callow workers in colonies in
which older workers were experimentally removed matured behaviorally in
one third the time of normal development and began foraging at an earlier age
than workers in unmanipulated control colonies (69). This type of socially
mediated behavioral development appears to be common in ants (59).
Flexibility in temporal polyethism appears to represent a major mechanism of
social compensation of forager number in response to short-term ecological
nuctuations.
Studies of trail system organization in Formica spp. have illustrated the
role of age polyethism in the development of foraging behavior (15, 87).
Foragers captured and marked in autumn and recaptured in the spring retained
204 TRANIELLO

the use of their initial route through a period of winter inactivity. These
topographic traditions result in a transfer of spatial foraging infonnation
through several worker generations. ROlltes have a spatiotemporal polyethic
structure; mature (pioneer) foragers head the foraging column, whereas youn­
ger workers (recruits) are found close to the nest. The behavior of young
foragers gradually changes to that of mature foragers, as evidenced from
changes in their location along a foraging trail.
Because foraging behavior is often coordinated through chemical com­
munication, maturational, social, or perceptual abilities may be associated
with the onset of foraging. Callow army ants (N. nigrescens) travel along
chemical trails with other colony members during emigration on the first
nomadic day but do not begin foraging (raiding) until they are 3-7 days old.
The absence of foraging behavior is not due to the sensory inability of
callows, however, as these immature workers follow artificial trails as well as
adults; this suggests that other social stimuli affect the development of
foraging (02). That callows remain in the nest while mature workers conduct
raids seems to be due to the absence of mechanical stimuli that are part of the
mass-recruitment communication signals that accompany emigration and are
required by callows to induce trail following (103).

LEARNING AND FORAGING BEHAVIOR

The unpredictability of the resource environment of a colony may require that

foraging be fine-tuned through experience-based change in the behavior of
individual workers to enhance efficiency. Simple and complex types of
learning have been described in ants, which may reflect selection for learning
traits that have evolved in response to dietary habits and foraging ecology
under the constraints of life span, sociality, and caste and the temporal,
spatial, and size variability of food resources. Recent studies of ant learning
have emphasized either ecology (81) or theoretical modeling (27, 28); addi­
tional empirical studies on the natural history of ant learning will be useful in
defining the limits of individual flexibility and measuring forager competence
(44). Veromessor pergandei and P. rugosus foragers develop preferences for
harvesting the seeds of certain plants, which results in an individual foraging
specialization that may persist for up to two weeks or longer (81). Learning
rate and memory retention may be correlated with spatial variability in seed
species in these ants (R. Johnson, personal communication). Similar forager
specializations appear to occur in Chelaner spp. (25). The mechanism of
specialization may be a learning process that permits a forager to identify
qualitative properties of seeds and to narrow search image quickly to fix on a
specific foraging task. This specialization may increase the net rate of energy
returned to the colony. Learning may also have a role in the development of
 ANT FORAGING STRATEGIES 205

foraging in other species. In omnivorous species such as Formica spp. and

Cataglyphis spp., workers tend homopterans or lick plants to collect carbo­
hydrate secretions and also forage for arthropod prey (111, 112, 119). These
two resources differ in their spatial and temporal patterns of distribution and
therefore provide different patterns of reward. Workers of C. bicolor tend to
specialize on collecting either type of food (39). In F. schaufussi, the ability
of workers to modify their search behavior seems to depend on a food
resource-dependent learning process. Local search duration is more easily
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.

conditioned by carbohydrate food rewards (sucrose) than by protein (freshly

killed insects) (111, 112). The conditioning of search duration by carbo­
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org

hydrate food seems to be independent of resource quality. Repeated rewards

of insect prey do not produce a more accurate search pattern, nor do they
result in increased search effort. Search accuracy is greater after a reward of a
single crop load of 1-M sucrose than after a reward of insect prey. However,
for a given food type, there is no significant change in search accuracy over
four or eight consecutive rewards (J. F. A. Traniello & A. 1. Kozol, un­
published). Further studies are needed to determine if foraging specializations
develop through resource-related learning processes. Resource-related learn­
ing processes may also be involved in the persistence of route fidelity and
memory shown in Formica spp. (18, 88). At the colony level, learning may
underlie competitor recognition systems and may mediate community in­
teractions (13).

CONCLUSIONS

The principal ecological determinants of ant foraging strategy are the distribu­
tion of food resources in size, time, space, and quality; competition with
sympatric ant species; and predation. Because a colony is generally sessile,
the resource and competitive environment is in essence defined by its loca­
tion. A colony must therefore spatially pattern foraging to harvest food
efficiently and minimize competition. This is accomplished through in­
dividual and colony-wide foraging.
Elements of social organization are linked to virtually every aspect of
foraging strategy. Although patterns of physical and temporal caste evolution
have been shown to be integral components of some foraging systems, more
sensitive empirical tests are required to provide detailed explanations of the
origin and ecological significance of forager size and age-related behavior.
Similarly, forager functional morphology and biomechanical characteristics
associated with loading capacity should be examined to describe precisely the
role of form as well as size and behavior in ant foraging ecology. The overall
conceptualization of ant foraging strategy should therefore encompass theory
and analyses on subjects ranging from the major elements of colony organiza-
 206 TRANIELLO

tion such as caste and division of labor to the time and energy budgets of
foragers. Although in this review I have suggested a components approach,
foraging systems must be analyzed as whole entities. The pitfalls of extreme
adaptationist reasoning that might develop from fragmenting the system
should and can be avoided through an emphasis on social integration. A
prudent application of foraging models, theories on caste evolution and
ecology, and basic knowledge of natural history and social regulatory mech­
anisms can provide the necessary theoretical and empirical framework. Final­
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.

ly, it will be possible to evaluate ant foraging strategy fully when the
energetics of foraging and the caloric and nutritional benefits of selected
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org

resources are quantitatively measured and used to estimate the conversion of

food intake into alates. This analysis will remove the inaccuracies of assump­
tions of the energetic costs of foraging and will permit colony foraging
economics to be quantified in terms of reproductive output.

ACKNOWLEDGMENTS

I thank many colleagues for their reprints and unpublished manuscripts, and
regret that a lack of space prohibited me from including the results of a larger
number of studies. The manuscript benefited from critical readings by Samuel
Beshers, Michelle Scott, and Steve Rissing. I gratefully acknowledge the
support of the National Science Foundation and the Whitehall Foundation.

Literature Cited
f. A. 1981.
1. Adams, E. A . , Traniello, J.
Chemical interference competition by
Monomorium minimum. Oecologia 5 1 :
270-75
2. Andersen, A. N. 1986. Patterns of ant
community organization in mesic south­
eastern Australia. Aust. J. Ecol. 11:87-
97
3. Bartholomew, O. A . , Lighton, J. R. B . ,
Feener, D . H . 1988. Energetics o f trail
running, load carriage, and emigration
in the column-raiding army ant Eciton
hamatum. Physiol. Zool. 61:57-68
4. Bernstein, R. 1974. Seasonal food abun­
dance and foraging activity in some des­
ertants. Am. Nat. 108:490-98
5. Bernstein, R. 1975. Foraging strategies
of ants in response to variable food den­
sity. Ecology 56:213-19
6. Bowers, M. A . , Porter, S. D. 1981.
Effect of foraging distance on water con­
tent of substrates harvested by AUa co­
lumbica (Guerin). Ecology 62:273-75
7. Breise, D. T. 1982. Patterning of re­
sources amongst seed-harvesting ants in
an ant community in semi-arid Aus­
tralia. Aust. J. Ecol. 7:299-307
8. Breise, D. T . , Macauley, B. J. 1981:
Food collection within an ant communi­
ty in semi-arid Australia, with special
reference to seed harvesters. Aust. J.
Ecol. 6:1-19
9. Brian, M. V. 1978. Production Ecology
of Ants and Termites. Cambridge, M ass:
Cambridge Univ. Press
1 0 . Brian, M . V. 1983. Sociallnsects: Ecol­
ogy and Behavioural Biology . London:
Ch apman & Hall
11. Buckley. R. C. 1982. Ant-Plant In­
teractions in Australia. The Hague: Junk
12. Burton, J. L . , Franks, N. R. 1985. The
foraging ecology of the army ant Eciton
rapax: an ergonomic enigma? Ecol. En­
tomo!' 10:131-41
13. Carlin , N. F . , Johnston, A. 1 984.
Learned enemy specification in the de­
fense recruitment system of an ant.
Naturwissenschaften 71: 156--57
14. Carroll, C. R . , Janzen, D. H. 1973.
Ecology of foraging by ants. Ann. Rev.
Ecol. Syst. 4:231-57
15. Cherix, D. 1987. Relation between diet
and polyethism in Formica culonies. See
Ref. 76, pp. 93-115
 ANT FORAGING STRATEGIES
1 6 . Chew, R. M. , Chew, A. E. 1 980. Body
size as a detenninant of small-scale dis­
tribution of ants in evergreen woodland
in southeastern Arizona. lnsectes Soc.
27: 1 89-20
17. Chew, R. M . , DeVita, 1. 1 980. Forag­
ing characteristics of a desert ant assem­
blage: functional morphology and spe­
cies separation. J. Arid Environ . 3:75-
83
1 8 . Cosens, D., Toussaint, N. 1985. An ex­
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.
perimental study of the foraging strategy
of the wood ant Formica aquilonia .
Anim. Behav. 33 :54 1-52
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org
1 9 . Crawford, D. L. , Rissing, S. W. 1983 .
Regulation of recruitment by individual
scouts in Formica oreas Wheeler (Hy­
menoptera; Formicidae). lnsectes Soc.
30: 1 77-83
20. Culver, D. 1974. Species packing in
Caribbean and north temperate ant com­
munities . Ecology 55:974-88
2 1 . Davidson, D. W. 1 977. Species diver­
sity and community organization in des­
ert seed-eating ants. Ecology 5 8 : 7 1 1-24
22. Davidson, D. W. 1 977. Foraging ecolo­
gy and community organization in seed­
eating ants. Ecology 58:725-37
23. Davidson, D. W. 1978. Experimental
tests of the optimal diet in two social
insects. Behav. Eeol. Sociobiol. 4:35-41
24. Davidson, D. W . 1978. Size variability
in the worker caste of a social insect
(Veromessor pergandei Mayr) as a func­
tion of the competitive environment.
Am. Nat. 1 1 2:523-32
25 . Davison, E. A. 1982. Seed utilization by
harvester ants. See Ref. 1 1 , pp. 1-6
26. Davison, E. A. 1987. Respiration and
energy flow in two Australian species of
desert harvester ants, Chelaner roth­
steini and Chelaner whitei. J. Arid. En­
viron . 1 2:61-82
27. Deneubourg , 1 . L. 1985. The role of
errors and memory in the efficiency of
ant foraging. Ann. Soc. R . Zool. Belg.
1 1 5 : 1 09-10
2lf. Deneubourg, J. L . , Goss, S. , Pasteels,
1. M. , Fresneau, D. , Lachaud, 1 . P.
1987. Self-organizing mechanisms in
ant societies (II): Learning in foraging
and division of labor. See Ref. 76, pp.
1 77-96
29. DeVita, J. 1979. Mechanisms of in­
terference and foraging among colonies
of the harvester ant Pogonomyrmex cali­
fornicus in the Mojave Desert. Ecology
60:729-37
30. Downing, H. 1978. Foraging and mi­
gratory behavior of the ponerine ant
Tennitopone laevigata. BA thesis.
Smith Coil. , Northampton, Mass. 1 2 1
pp.
207
3 1 . Fellers, J. H. 1987. Interference and ex­
ploitation in a guild of woodland ants.
Ecology 68: 1466-78
32. Fewell, J. 1988. Energetic and time
costs of foraging in harvester ants,
Pogonomyrmex occidentalis. Behav.
Ecol. Sociobiol. 22:401-8
33. Franks, N. R. 1985. Reproduction,
foraging efficiency, and worker poly­
morphism in army ants. In Experimental
Behavioral Ecology and Sociobiology,
ed. B . Holldobler, M. Lindauer, pp. 81-
108. Stuttgart: Fischer
34. Franks, N. R. 1986. Teams in social
insects: group retrieval of prey by army
ants (Eciton . burchelli, Hymenoptera:
Formicidae). Behav . Ecol. Sociobiol.
1 8:425-29
35. Franks, N. R. , Fletcher, C. A. 1983.
Spatial patterns in anny ant foraging and
migration: Eciton burchelli on Barro
Colorado Island, Panama. Behav. Eco!.
Sociobiol. 1 2:26 1-70
36. Fresneau, D. 1985. Individual foraging
and path fidelity in a ponerine ant. In­
sectes Soc. 32: 1 09- 1 6
3 7 . Gentry, 1 . B . 1974. Response to preda­
tion by colonies of the Florida harvester
ant Pogonomyrmex badius. Ecology
5 5 : 1328-38
38. Hansen, S. E. 1978. Resource utiliza­
tion and coexistence of three species of
Pogonomyrmex ants in an upper Sonoran
grassland community. Oeeologia 35:
105-17
39. Harkness, M . L. R . , Harkness, R. D.
1 976. Functional differences between
individual ants Cataglyphis bicolor. J.
Physiol. 25: 1 24-25
40. Harkness, R. D. 1978. The speed of
walking of Cataglyphis bicolor (F.)
(Hym . , Formicidae). Entomol. Mon.
Mag . 1 1 4:203-1 0
4 1 . Harkness, R . D. , Maroudas, N. G.
1 9 8 5 . Central place foraging by an ant
(Caraglyphis bicolor Fab.): a model
of searching. Anim. Behav. 33:916-
28
42. Harrison , 1 . S .. Gentry, 1 . B . 1 98 1 .
Foraging pattern, colony distribution,
and foraging range of the Florida harves­
ter ant, Pogonomyrmex badius. Ecology
62: 1467-73
43. Heinrich, B. 1978. The economics of
insect sociality. In Behavioural Ecology,
ed. 1. Krebs , N. Davies, pp. 97- 1 2 8 .
Sunderland, Mass: Sinauer
44. Herbers , 1. M. 1 98 1 . Reliability theory
and foraging by ants. J. Theor. Bioi.
89: 1 75-89
45. Herbers, 1 . M. 1985 . Seasonal structur­
ing of a north temperate ant community.
lnsectes Soc. 32:224-40
 208 TRANIELLO
46. Hespenheide, H. A. 1973. Ecological
inferences from morphological data.
Ann. Rev. Ecol. Syst. 4 : 2 1 3-3 0
47. Hespenheide, H. A. 1975. Prey charac­
teristics and predator niche width. In
Ecology and Evolution of Communities,
ed. M. L. Cody, J. M. Diamond, pp.
1 5 8-80. Cambridge, Mass: Belknap
48. Holder Bai ley , K . , Polis, G. A. 1987.
Optimal and central place foraging
theory applied to a de�ert harvester ant,
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.
Pogonomyrmex californicus. Oecologia
72:440-48
49. Hiilldobler, B. 1 976. Recruitment be­
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org
havior, home range orientation and ter­
ritoriality in harvester ants, Pogono­
myrmex. Behav. Ecol. Sociobiol. 1 :3-
44
50. Hiilldobler, B . 1 984. Evolution of insect
communication. In Insect Communica­
tion, ed. T. Lewis, pp. 349-77 . London:
Academic
5 1 . Hiilldobler, B . , Lumsden, C 1 980. Ter­
ritorial strategies in ants. Science 2 1 0:
732-39
5 2 . Hiilldobler, B . , Moglich, M. 1 980. The
foraging system of Pheidole militicida
(Hymenoptera : Formicidae) . Insectes
Soc. 27:237-64
5 3 . H611dobler, B . , Stanton, R. C . , Markl ,
H. 1 97 8 . Recruitment and food-retriev­
ing behavior in Novomessor (Formici­
dae: Hymenoptera) . 1. Chemical signals.
Behav. Eco/. Sociobiol. 4: 1 63-8 1
54. H61 1dobler , B . , Traniello, J. F. A. 1980.
The pyg idi al gland and chemical recruit­
ment communication in Pachycondyla
( = Termitopone) laevigata . J. Chem.
Eco!. 6:883-93
55. Howard, 1. 1. 1 987. Leafcutting ant diet
selection: the role of nutrients, water,
and secondary che mistry . Ecology 68:
503- 1 5
56. Hunt, J . H . 1983. Foraging and mor­
phology in ants: the role of vertebrate
predators as agents of natural selection.
See Ref. 57, 2:83-103
57. Jaisson, P. 1983. Social Insects in the
Tropics, Vol. I , 2. Paris: Presses Univ.
Paris XIII
5 8 . Johnson, L. K . , Hubbell, S. P . , Feener,
D . H. 1987. Defense of food supply by
eusocial colonies. Am. Zool. 27:347-58
59. Lenoir, A. 1 987 . Factors determining
polyethism in social insects. See Ref.
76, pp. 2 1 9-40
60. Leonard, 1. G . , Herbers, 1. M. 1986.
Foraging tempo in two woodland ant
species. Anim. Behav. 34: 1 1 72-8 1
6 1 . Levings, S. C , Franks, N. R. 1982.
Patterns of nest dispersion in a tropical
ground ant community. Ecology 63:
33 8-44
62, Levings, S. C . , Traniello, J. F. A, 1982,
Territoriality, nest dispersion, and com­
munity structure in ants. Psyche 88:265-
319
63 . Lighton, 1 . R . B . , Bartholomew, G . A. ,
Feener, D. H. 1987. Energetics of loco­
motion and load carriage and a model of
the energy cost of foraging in the leaf­
cutting ant Atta colombica Guer. Physi-
01. Zool. 60:524-37
64. Longhurst, C . , Johnson, R. A . , Wood,
T. G. 1978. Predation by Megaponera
foetens (Fabr. ) (Hymenoptera: Formici­
dae) on termites in the Nigerian southern
guinea savanna. Oecologia 32: 1 0 1-7
65 . Lynch, J. F . , Balinksy, E. C , Vah, S .
G . 1980: Foraging patterns i n three sym­
patrie forest ant species, Prenolepis im­
puris, Paratrechina melanderi, and
Aphaenogaster rudis (Hymenoptera:
Formicidae). Ecol. Entomol. 5:353-7 1
66. MacKay, W. P. 1982. The effect of pre­
dation of western widow spiders Lat­
rodectus hesperus (Araneae:Theridiiae)
on harvester ants Pogonomyrmex rugo­
sus (Hymenoptera:Formicidae). Oecolo­
gia 53 :406-1 1
67. MacKay, W. P. 1 985. A comparison of
the energy budgets of three species of
Pogonomyrmex harvester ants (Hyme­
noptera: Formicidae). Oecologia 66:
484-94
68. Maschwitz, U . , Schonegge , P. 1983.
Forage communication, nest moving
recruitment and prey specialization in
the oriental ponerine Leprogenys chinen­
sis . . Oecologia 57: 1 75-82
69. McDonald, P . , Topoff, H . 1 985. Social
regulation of behavioral development in
the ant Novomessor albisetosus (Mayr).
J. Camp. Psycho/' 99:3- 1 4
70. M i renda , J. T . , Eakin s, D . G . , Gravelle ,
K . , Topoff, H. 1980. Predatory be­
havior and prey selection by army ants in
a desert-grassland habitat. Behav. Eco/.
Sociobiol. 7: 1 1 9-27
71. Mirenda, J. T . , Topoff, H. 1980.
Nomadic behavior of army ants in a des­
ert-grassland habitat . Behav. Ecol. So­
ciobiol. 7: 1 29-35
72. Morton , S. R. 1982. Granivory in the
Australian arid zone: diversity of harves­
ter ants and structure of their communi­
ties. In Evolution of the Flora and
Fauna of Arid Australia, ed. W. R. Bar­
ker, P. J. M. Greenslade, pp. 257-62.
Adelaide, Australia: Peacock
73. Nielsen, M. G . , Jensen, T. F . , Holm­
Jensen, 1 . 1982 . Effect of load carriage
on the respiratory metabolism of running
worker ants of Camponotus herculeanus
(Formicidae). Oikos 39: 1 37-42
74. Nonacs, P . , Dill, L. M. 1988. Foraging

responses of the ant Lasius pallitarsus to
food sources with associated mortality .
Insectes Soc. In press
75. Oster, G . , Wilson, E. O. 1978. Caste
and Ecology in the Social Insects.
Princeton , N1: Princeton Univ. Press
76. Pasteels, J. M . , Deneubourg, 1. L.
1 987. From Individual to Collective Be­
haviour in Social Insects. Basel: Birk­
hauser
77. Peakin, G. J . , Josens, G. 1 978. Respira­
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.
tion and energy flow. See Ref. 9, pp.
1 1 1-63
78. Porter, S.D . , Jorgenson, C. D . 1 98 ! .
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org
Foragers o f the ant, Pogonomyrmex
owyheei: a disposable caste? Behav .
Ecol. Sociobiol. 9:247-56
79. Rettenmeyer, C. ' W . , Chadab-Crepet ,
R . , Naumann, M. G . , Morales, L.
1983. Comparative foraging by neotro­
pical anny ants. See Ref. 57, 2:59-73
80. Reyes Lopez, J. L. 1987. Optimal forag­
ing in seed harvester ants: computer­
aided simulation. Ecology 68:1630--33
8 1 . Rissing, S. W. 1 98 1 . Foraging spe­
cializations of individual seed-harvester
ants. B ehav . Ecol. Sociobiol. 9: 1 49-
52
82. Rissing, S. W. 1982. Foraging velocity
of seed-harvester ants, Veromessor per­
ga nde i (Bymenoptera: Fonnicidae). En
viron. Entomol. 1 1 :905-7
83. Rissing, S. W. 1987. Annual cycles in
worker size of the seed-harvester ant
Veromessor pergandei (Hymenoptera:
Formicidae) . Behav. Ecol. Sociobiol.
20: 1 1 7-24
B . 1 984.
84. Rissing, S . W. , Pollock, G.
Worker size variability and foraging
efficiency in Veromessor pergandei
(Hymenoptera: Formicidae). Rehav.
Eml. Sociobiol. 1 5 : 1 2 1 -26
85. Rissing, S. W . , Wheeler, J. 1 976. Fo­
raging responses of Veromessor per­
gandei to changes in seed production.
Pan-Pac. Entomol. 52:63-72
86. Rockwood, L . , Hubbell, S. P. 1987.
Host-plant selection, diet diversity. and
optimal foraging in a tropical leafcutting
ant. Oecologia 74:55-61
87. Rosengren, R. 1 977. Foraging strategy
of wood ants, Formica rufa group. Pan
I . Age polyethism and topographic
traditions. Acta Zool. Fenn. 149: 1 3 0
- ant Neivamyrmex nigrescens: impor­
88. Rosengren , R . , Fortelius, W. 1986. Or­
streue in foraging ants of the Formica
rufa group--hierarchy of orienting cues
and long-term memory . Insectes Soc.
33:306-37
89. Rudolph, S. G Loudon, C. 1986. Load
..
size selection by foraging leaf-cutter ants
(Aua cephalotes). Ecol. Entomol. 1 1:
40 1 - 10
ANT FORAGING STRATEGIES 209
90. Ryti, R. T. , Case, T. J. 1 986 . Over­
dispersion of ant colonies: a test of hy­
potheses. Oecologia 69:446-53
9 1 . Schmid-Hempel, P. 1984. Individually
different foraging methods in the desert
ant Cataglyphis bicolor (Hymenoptera,
Formicidae). Behav. Ecol. Sociobiol.
14:263-7 1
92. Schmid-Hempel , P. 1 987. Foraging
characteristics of the desert ant Cata­
glyphis. See Ref. 76, pp. 43-61
93. Schmid-Hempel, P . , Schmid-Hempel,
R . 1984. Life duration and turnover of
foragers in the ant Cataglyphis bicolor
(Hymenoptera, Formicidac). lnsectes
Soc . 3 1 :345-60
94. Shepherd, J. D. 1 982. Trunk trails and
the searching strategy of a leaf-cutter ant
A lta colombica . Behav. Ecol. Sociobiol.
1 1 :77-84
95. Shepherd , J. D. 1985 . Adjusting forag­
ing effort to resources in adjacent colo­
nies of the leaf,cutter ant, Atta col­
ombica. Biotropica 1 7:245-52
96. Sorensen , A. A . , Busch, T. M . , Vinson,
S. B . 1983. Behaviour of worker sub­
castes in the fire ant, Solenopsis invicta,
in response to proteinaceous food. Phys­
iol. Entomol. 8:83-92
97 . Stephens, D. W . , Krebs, 1. R. 1986.
­ Foraging Theory . Princeton, NJ: Prince­
ton Univ. Press
98. Stradling, D. J. 1978. Food and feeding
habits of ants. See Ref. 9, pp. 8 1 -
1 06
99. Sudd, 1. H. 1 987. Individual behavior
and mixed diet strategy in ants. See Ref.
76. pp. 8 1-92
100. Taylor. F. 1976. Foraging behavior in
ants: experiments with two species of
myrmecine ants. Behav. Eco!. Socio­
bioi. 2: 1 47-68
1 0 1 . Taylor, F. 1 978. Foraging behavior of
ants: theoretical considerations . J.
Theor. Bioi. 7 1 :54 1-65
1 02. Topoff, H . , Boshes, N . , Trakimas, W.
1972. A comparison of trail following
between callow and adult workers of the
army ant (Neivamyrmex nigrescens)
(Formicidae: Dorylinae). Anim. Behav.
20:36 1 -66
1 0 3 . Topoff, H . , Mirenda 1. 1978. Precocial
,
behaviour of callow workers of the army
tance of stimulation by adults during
mass recruitment. Anim. Behav. 26:
698-706
104. Topoff, H . , Mirenda, J. 1980. Army
ants do not eat and run: influence of food
supply on emigration behaviour in
Neivamyrmex nigrescells. Anim. Behav.
28: 1040--45
105. Topoff, B . , Mirenda, J. 1 980 . Army
 210 TRANIELLO
ants on the move: relation between food
supply and emigration frequency. Sci­
ence 207 : 1 099-1 00
106. Traniello, 1. F. A. 1 977 . Recruitment
behavior, orientation, and the organiza­
tion of foraging in the carpenter ant
Camponotus pennsylvanicus DeGeer.
Behav. Ecol. Sociobiol. 2:61-79
1 07 . Traniello, J. F. A. 1978. Caste in a
primitive ant: absence of age polyethism
in Ambylopone. Science 202:770-72
by Indiana University - Purdue University Indianapolis - IUPUI on 09/29/12. For personal use only.
lOS. Traniello, J. F. A. 1 9S0. Colony
specificity in the trail pheromone of an
ant. Naturwissenschaften 67;360
Annu. Rev. Entomol. 1989.34:191-210. Downloaded from www.annualreviews.org
1 09. Traniello, J . F. A . 1983. Social organ­
ization and foraging success in Lasius
neoniger (Hymenoptera; Formicidae);
behavioral and ecological aspects of
recruitment communication. Oecologia
59;94- 1 00
1 1 0. Traniello, J. F. A. 1 987. Comparative
foraging ecology of north temperate
ants: the role of worker size and coop­
erative foraging in prey selection. In­
sectes Soc. 34: 1 1 8-30
1 1 1. Traniello, J. F. A. 1 987 . Individual and
social modification of behavior in re­
sponse to environmental factors in ants.
See Ref. 76, pp. 63-80
1 12. Traniello, 1. F. A. 1 988. Variation in
foraging behavior among workers of the
ant Formica schaufussi: ecological
correlates of search behavior and the
modification of search pattern. In In­
terindividual Behavioral Variability in
Social Insects, ed . R. L. Jeanne : pp.
9 1- 1 1 2 . Boulder, Colo; Westview
1 1 3 . Traniello, J. F. A. 1 988. Chemical trail
systems, orientation, and territoriality in
the ant Lasius neoniger. i. lnsect Behav.
Tn press
1 1 4. Traniello, J. F. A . , Fujita, M. S . , Bo­
wen, R. V. 1 984 . Ant foraging be­
havior: Ambient temperature influences
prey selection. Behav. Ecol. Sociobiol.
1 5 :65-68
1 15. Traniello, J. F. A . , Levings, S. C. 1 986.
Intra- and intercolony patterns of nest
dispersion in the ant Lasius neoniger:
correlations with territoriality and forag­
ing ecology. Oecologia 69:4 1 3- 1 9
1 1 6. Waller, D. A . 1 982. Leaf-cutting ants
and avoided plants; defenses against Alta
texana attack. Oecologia 52:400-3
1 1 7. Waller, D. A. 1 982 . Leaf-cutting ants
and live oak; the role of leaf toughness in
seasonal intraspecific host choice . En­
tomol. Exp . Appl. 32; 1 45-50
1 1 8. Wehner, R. 1 987. Spatial organization
of foraging behavior in individually
searching desert ants, Cataglyphis (Sa­
hara Desert) and Ocymyrmex (Namib
Desert). See Ref. 76, pp. 1 5-42
1 1 9. Wehner, R . , Schmid-Hempel , P . , Hark­
ness, R. D. 1 983. Foraging Strategies in
Individually Searching Ants Cataglyphis
bicolor. Stuttgart: Fischer
1 20 . Whitford , W. G. 1 978. Foraging by
seed-harvester ants. See Ref. 9, pp.
\ 07- 1 1 0
121. Whitford , W. G . , Bryant, M . 1 979. Be­
havior of a predator and its prey; the
homed lizard (Phrynosoma cornutum)
and harvester ants (Pogonomyrmex
spp.). Ecology 60:686-94
1 22 . Wilson, D. S. 1 975 . The adequacy of
body size as a niche indicator. Am. Nat.
\ 09:769-84