Arthropod Predation by a Specialist Seed Predator, the Golden-backed Uacari (Cacajao melanocephalus ouakary, Pitheciidae) in Brazilian Amazonia

Morphological adaptations related to food processing generally reflect those elements of the diet that represent the greatest biomechanical challenge or that numerically dominate the diet. However, in periods of the annual cycle when the availability of such foods is low, items to which a species has low apparent morphological adaptation may be included in the diet. Here we test the responses of a diet-specialist primate to limitations in the supply of the resource it is specialized to exploit. Uacaris are primarily predators of immature seeds, in seasonally flooded forests in Amazonian Brazil, and have dental specializations to open hard-shelled fruits. We investigated the importance of arthropods in the diet of golden-backed uacaris (Cacajao melanocephalus ouakary), examining their seasonal importance in the uacari diet, and the ways C. m. ouakary used to access them. Using scan and ad libitum sampling of feeding and phenology from botanical study plots to assess fruit availability, we conducted an 18-mo study in Jaú National Park, Amazonas State, Brazil. We recorded arthropod predation 298 times, with Cacajao melanocephalus ouakary feeding on 26 invertebrate taxa in ≥11 families and 9 different orders. Uacaris extracted wood-boring beetles dentally from rotting wood and smaller larvae from twigs, stems, and petioles, but this food class did not predominate. This food class (encapsulated foods) constituted 23.4 % of the arthropod records. The majority of arthropod food items were either manually removed from substrates (ants, beetle larvae, caterpillars, fulgorid bugs, grasshoppers, mayflies, spiders, termites, wasps, and a whip-scorpion) or plucked from the air (volant Lepidoptera). Uacaris appeared to avoid toxic caterpillars. Insectivory was most frequent when fruit and seeds were least available. Arthropods seem to be seasonally important to this primate, supplementing or making up for shortfalls in the hard fruits and immature seeds for which uacaris have highly developed dental, and possibly intestinal, adaptations.     

Diet,Habitat Selection and Natural History of <Emphasis Type="Italic">Cacajao melanocephalus ouakary</Emphasis> in Jaú National Park,Brazil<Superscript>1</Superscript>

We studied golden-backed uacaris, (Cacajao melanocephalus ouakary) in the late-wet season of 1999 and in the mid-dry season of 2000 in Jaú National Park (JNP), Brazil. Additional observations were taken by other scientists working in JNP. Group counts range from 2 to 30 in the wet season and from 1 to > 100 in the dry. In the wet season, we recorded uacaris only in flooded forests (igapó). During the dry season they were in igapó, campinarana, and terra firme forests. We supplemented observations via interviews with local informants on seasonal use of the forest types and of particular small patches of different habitat types that exhibit high temporal patchiness in fruit availability. We recorded a total of 13 food plant species (dry season, 4; flooded season, 9), and a further 55 were provided by interviews. We noted the physical dimensions and hardness of 26 fruit species. Despite accepted specializations of the uakaris on hard unripe fruit, we recorded incidents of insectivory and folivory. Patterns of food availability, and therefore the seasonal habitat-use patterns by golden-backed uacaris, appear to be more complex than previously realized.The authors dedicate this paper to the memory of J. Marcio Ayres, the founding father of uacari field biology.     

Self-selection and utilization efficiency of Heliothis zea larvae feeding on seeds and the walls of snap bean pods

First instar Heliothis zea larvae tended to feed on the leaves of snap beans, but later instars preferred to burrow in the pods. Fifth instars offered only leaves grew poorly because they ate them in small quantities, presumably because of deterrency. The utilization efficiency and growth of fifth instars fed pulp was slightly but significantly inferior to that of seed-fed larvae but was greatly superior to that of leaf-fed larvae. Since the dry matter content of pulp is less than one-third that of seeds, larvae allowed to eat only pulp had abnormally large fresh weight intakes and devoted about three times as much time to feeding as did seed-fed larvae. Larvae offered a mix of seeds and pulp ate both and had a total fresh weight food intake somewhat greater than that of seed-fed larvae but much less than that of pulp-fed larvae. Eating both seeds and pulp offers no obvious nutritional advantage but presumably offers a significant ecological advantage. It is argued that the number of foraging trip a larva must make to discover pods is minimized by feeding on a mix, thus reducing exposure to mortality factors such as predation.     

Temporal patterns of seed use and availability in a guild of desert ants

ABSTRACT.

• 1 Temporal patterns of seed use were studied from late winter to autumn in three species of seed-harvesting ants in the Sonoran Desert. Measures of effective foraging activity, dietary niche breadth and dietary niche overlaps were obtained each month and were tested for correlation with estimates of the available seed resource.
• 2 Seeds were the only numerically important type of food in the diets of all species.
• 3 The ants partitioned the resource according to both seed species and seed size, although there was considerable overlap.
• 4 Pheidole xerophila had the smallest forager body size and is a specialist on small seeds because it harvested them in greater proportion than their rank in the soils and expanded its diet to larger seeds only when the abundance of small seeds declined.
• 5 When the abundance of the small seeds of Bouteloua barbata decreased, the middle-sized ant, Veromessor pergandei, showed a decrease in foraging activity, increase in niche breadth, and a decrease in overlap with P.xerophila.
• 6 Seed size preferences of V.pergandei did not vary seasonally, except that during the month of highest seed abundance, V.pergandei showed no size preference.
• 7 Pogonomyrmex rugosus was the largest ant; it preferred larger seeds and was inactive when small seeds were most abundant. Seasonal foraging activity and niche parameters were random in relation to seed abundance.
• 8 We suggest that nocturnal foraging by P.rugosus during the summer months was a response to interference with diurnal foraging by either predation frorn horned lizards or competition from V.pergandei.
• 9 Seasonal abundance of small seeds explains most of the seasonal foraging patterns of P.xerophila and V.pergandei. The summertime abundance of larger seeds during years of adequate precipitation may account for the seasonal activity patterns of P.rugosus.

     

Individual foraging components of harvester ants: movement patterns and seed patch fidelity

Summary We investigated individual foraging components of the western harvester ant,Pogonomyrmex occidentalis, in the native seed background of a shrub-steppe environment. Our study identified factors affecting foraging movements and seed selection by individual ants. Some assumptions and predictions of central-place foraging theory and a correlated random walk were evaluated for individual foragers. Results showed that ant size was only weakly correlated with the seed sizes harvested; seed size was a more important constraint than a predictor of seed selection. Individual ants spent more time in localized search behavior than traveling between search areas and nests.P. occidentalis foragers encountered seeds randomly with respect to time, and handled a mean of 1.7 seeds/trip. A correlation of increased search effort with greater travel distances was consistent with central-place foraging theory but, contrary to it, search and travel effort were not associated with energetic reward.Individual ants exhibited fidelity in both search site and native seed species. Spatial analyses of foraging movements showed a highly oriented travel path while running, and an area-restricted path while searching. Searching ants moved in a manner consistent with a correlated random walk. The deterministic component of patch fidelity and the stochastic component of search may override energetic foraging decisions in individualP. occidentalis ants.     

Seasonal variation in fleshy fruit use and seed dispersal by the Japanese black bear (<Emphasis Type="Italic">Ursus thibetanus japonicus</Emphasis>)

Fruit use by the Japanese black bear (Ursus thibetanus japonicus) and seed clumping in bear scat were studied in central Japan using fecal analyses. Between May and November 2003 and 2004, the life form and fruit size of plants consumed by bears and the species composition and intactness of seeds contained in scat were examined in five transects (approximately 10 km × 10 m) in broad-leaved deciduous forests. In 2003, scats with seeds were found only in the autumn, when fruiting trees and shrubs were abundant. In 2004, scats with seeds occurred intermittently from the summer, when fruiting plants were rare, up to the autumn. Yearly and seasonal variation in fruit use reflects the opportunistic foraging behavior of Japanese black bears. Seven of the nine plant species detected in scats had medium-sized fruits (6–15 mm width), whereas the other two species had relatively large fruits (20–100 mm width). In total, 14,492 seeds were detected, of which 97.6% were intact; the remainder were damaged. Intact seeds of one or two species were found in each scat. The number of intact seeds per scat ranged from 1 to 5476. Japanese black bears seldom digest ingested seeds, thereby contributing to the seed dispersal of their food plants, including species with fruits that are too large to be swallowed by frugivorous birds.     

Ethogram and Natural History of Golden-backed Uakaris (<Emphasis Type="Italic">Cacajao melanocephalus</Emphasis>)

We present an ethogram for golden-backed uakaris (Cacajao melanocephalus), based on observations in the field and on a captive individual. We also provide additional observations on the ecology of the wild animals. We studied 3 free-living groups (maximum counts of 5, 15, and 26 individuals) during two wet-seasons (March–July 2007 and January–June 2008) in the flooded igapó forest of Jaú National Park, Amazonas, Brazil. The groups lived in close proximity but never mixed, because river channels separated them. Groups showed fission-fusion behavior, subgroup sizes varied within groups, and we observed 13 different subgroup compositions. The areas used by the groups were ca. 0.82, 2.35, and 2.45 km². We defined a total of 9 behavioral categories. In the wild, the amount of time allocated to the behaviors traveling and foraging/feeding differed between months, but we found no difference in the amount of time devoted to behavioral categories across 3 periods of the day, possibly as a result of the patchy and unpredictable distribution or availability of food patches. Further, the activity patterns varied among groups, perhaps as a reflection of the different group sizes and compositions and different range sizes. We recorded 34 feeding items for wild golden-backed uakaris between January and June 2008, mostly fruits and seeds. No significant variation in the number of different food types occurred across months. We recorded 6 primate species in the study areas. The uakaris neither mobbed nor fled from any other primate species, with the exception of white-fronted capuchin monkeys (Cebus albifrons). In addition, golden-backed uakaris sometimes fled when seeing giant otters (Pteroneura brasiliensis), perhaps indicating that the otters are potential predators of the monkeys. The ethogram and behavioral data provided here increase knowledge of the behavioral ecology of the elusive golden-backed uakari, and will facilitate future comparative studies.     

Optimal foraging: The responses of Peromyscus leucopus to experimental changes in processing time and hunger

Summary Food density, degree of hunger prior to foraging, and food processing difficulty were varied in order to determine their effects on the diet diversity of captive Peromyscus leucopus foraging for buried seeds. No relationship was found between diet diversity and food density. However, diet diversity exhibited a significantly positive relationship with hunger, and a significantly negative relationship with seed processing time. These results strongly support optimal foraging theory.     

Foraging behavior and morphology: seed selection in the harvester ant genus, Pogonomyrmex

Optimally foraging animals can be behaviorally or morphologically adapted to reduce the energetic and time costs of foraging. We studied the foraging behavior and morphology of three seed harvester ant species, Pogonomyrmex barbatus, P. desertorum, and P. occidentalis, to determine the importance of behavioral strategies and morphological features associated with load carriage in reducing the costs of foraging. We found that none of five morphological features we measured had a significant impact on seed selection. Also, body size did not influence running speed, an important variable in time costs of foraging. Temperature had the largest effect on running speed in these species. Our results show that these species have foraging strategies which minimize the time costs of traveling with seeds. We also describe a pattern where the running speed in individual-foraging species is less affected by increasing seed size than in trunk-trail foragers, when temperature and body mass are held constant. These results support previous work which showed that time costs are most important in seed selection for Pogonomyrmex, and suggest that central place foraging theory may need to accommodate variation in foraging strategy to more accurately predict optimal seed size selection in harvester ants. Received: 16 June 1997 / Accepted: 15 December 1997     

Fruit removal of Coprosma quadrifida (Rubiaceae) by birds in south-eastern Australia

Abstract Removal of the fleshy fruits of Coprosma quadrifida by birds was examined in relation to fruit crop size, neighbouring fruit crop size and time in the season in montane wet sclerophyll forest in south-eastern Australia. On average, 84% of individual fruit crops were removed. Frugivores removed fruit in proportion to its abundance, indicating that they continually assess availability rather than respond to a critical abundance. The size of neighbouring fruit crops did not influence fruit removal rates, suggesting that there is little competition among neighbours for dispersers. The ripe fruits of C. quadrifida were eaten by 14 bird species, comprising 50% of the bird species trapped. Silvereyes (Zosterops lateralis) were the most abundant consumers of fruits. With the exception of the parrot, Platycercus elegans, all birds defecated seeds intact and had no influence on seed viability.     

THE RELATIVE COSTS OF SELF- AND CROSS-FERTILIZED SEEDS IN IMPATIENS CAPENSIS (BALSAMINACEAE)

By using a generally applicable technique that involves monitoring the development and survivorship of flowers and seed capsules, I estimated the material and energetic costs of producing self- and cross-fertilized seeds in Impatiens capensis. All flowers and fruits on six plants were censused intensively for the two-month period of reproduction. Cleistogamous (selfing) flowers ripened seed in about 24 days, compared to about 36 days for the chasmogamous (outcrossing) flowers. In terms of dry weight, selfed seeds cost about two-thirds as much as outcrossed seeds: 12.4 versus 18.4 mg dry weight per seed. When adjusted to the currency of calories, and including an independent estimate of pollen and nectar production in outcrossing flowers, I estimate the costs to be about 65 and 135 calories per selfed or outcrossed seed. Sources of error include the accuracy of the estimates of flower and fruit weight, and possible differences among the developmental stages in respiratory costs. The cost discrepancy implies that outcrossed seeds should possess a countervailing fitness advantage large enough to offset their greater energetic cost.     

Fruit availability and chimpanzee party size at Kahuzi montane forest,Democratic Republic of Congo

The effect of fruit availability on chimpanzee party size was investigated in the montane forest of Kahuzi. Seasonal variation in both fruit availability and party size was examined. Fruit abundance per se does not affect chimpanzee party size. However, seasonality and distribution patterns of fruits are both determinant ecological factors that control the size of chimpanzee parties at Kahuzi. There was no correlation between fruit abundance and the spatial distribution of fruits. When fruits were clumped and available in large amounts for a long period, chimpanzee party size increased, or otherwise decreased when fruits were highly available for only a limited period. Tree species that produced only a small amount of ripe fruit throughout the year did not affect the foraging party size of chimpanzees. Temporal and spatial variability in fruit abundance seems to constrain grouping patterns of chimpanzees at Kahuzi more so than in other chimpanzee habitats previously described.     

Foraging behaviour of harvesting ants determines seed removal and dispersal

Harvesting ants can affect the regeneration of plants through at least two different processes: seed removal and seed dispersal. We analyse the role of different foraging strategies of ants on patterns of seed removal and dispersal by three Messor species with considerable differences in their foraging systems. Messor capitatus workers rarely leave the nest in well-formed columns, while the other two species form foraging trails, with M. bouvieri forming temporary trails and M. barbarus foraging on a stable system of permanent foraging trails. Overall seed intake of M. capitatus colonies is considerably less than that of the two group-foraging species. There are also differences in the size of seeds collected: M. barbarus and M. capitatus harvest similar amounts of large and small seeds, while M. bouvieri harvests small seeds more intensely than large ones, due to the smaller size of the worker caste. The three Messor species differ in the percent of seed dropping of the different seed type and in the seed dispersal distance. Moreover, M. bouvieri and M. capitatus redistributed dropped seeds preferentially in bare soil and low sparse vegetation habitats, while M. barbarus redistributed seeds mainly in the high vegetation habitat. These results show that the foraging systems of these harvesting ants determine different patterns of seed removal and dispersal and, thus, affect the abundance and redistribution of seeds in the area.     

Fruits foraging patterns and seed dispersal effect of frugivorous birds on Hippophae rhamnoides sinensis

Behaviors of 18 species of birds eating fruits of Hippophae rhamnoides spp. sinensis were observed from September 2003 to March 2004. Their foraging patterns were found to be very different and can be divided into five classes: (1) direct swallowing the fruits on crown of the shrubs and sometimes regurgitating seeds soon after; (2) carrying the fruits to their perching sites and swallowing; (3) pecking the fruits from the shrubs to the ground, eating pulp and seeds but leaving pericarp; (4) pecking through the pericarp, eating pulp and leaving pericarp and seeds; (5) pecking through the pericarp on the top of fruits, and only eating seeds. These foraging patterns have different effects on seed dispersal of H. rhamnoides spp. sinensis. The germination experiment of three groups of seeds (seeds from feces, dry fruits and extracted seeds from dry fruits) was carried out. Although ingestion processes of birds had some adverse effects on the seed germination of H. rhamnoides spp. sinensis, the seeds from feces still have a relatively higher germination ratio. H. rhamnoides spp. sinensis provides food to a variety of frugivorous birds, and the birds disperse its seeds. Thus, a mutually beneficial relationship between the bird and the seed is formed. __________ Translated from Chinese Journal of Ecology, 2005, 24(6): 635–638 [译自: 生态学杂志, 2005, 24(6): 635–638]     

Intensity of pre‐dispersal seed predation in the invasive legume Leucaena leucocephala is limited by the duration of pod retention

Abstract The intensity of seed predation the invasive tropical legume Leucaena leucocephala by the bruchid Acanthoscelides macropthalmus was investigated in south‐eastern Queensland, Australia. The number of seeds damaged by A. macropthalmus as a proportion of total seeds available was found to increase the longer the pods remained on the tree. Seed predation ranged from a mean of 10.75% of seeds on pods that remained on the plant for 1 month and increased to 53.54% for pods that remained of the plant for 4 months. The low bruchid populations at high pod densities results in ‘predator satiation’. However, pods dehisce over time and the proportion of pods available over time to the bruchid correspondingly declines. By the time bruchid densities build up, most pods have dehisced and the seeds consequently escape predation. As a result the number of seeds lost to bruchid damage increases only marginally over time. Despite the levels of seed predation observed over the course of the study, the number of seeds in the soil seedbank almost doubled over time increasing from 8.5 seeds m^?3 to 15.5 seeds m^?3 over a 4‐month period. Levels of seed predation and addition of seeds to the soil seedbank were not correlated. The taxonomic (subspecies) status and apparency of host plants as measured by plant and patch traits (average plant height, density of podding plants and patch size) did not influence levels of seed predation. Pre‐dispersal seed predation studies need to take into account the pod/seed retention behaviour of the plant. The ability of the bruchid to regulate the invasiveness of Leucaena through influencing its demography is likely to be diminished if the insect populations cannot increase rapidly enough to use the seeds before pod dehiscence.     

DEMOGRAPHIC ASPECTS OF FLOWER AND FRUIT PRODUCTION IN BEAN PLANTS,PHASEOLUS VULGARIS L.

In this paper we assess the relative contributions to total pod yield of cohorts of flowers of known age. Bean plant (Phaseolus vulgaris L.) growth was monitored and 'births' of individual flowers were recorded and their fates followed. For every cohort of flowers an array of yield measures was calculated, including the mean and total cohort dry weight of fruits, and the numbers of viable and aborted seeds and completely undeveloped ovules. When mature, pods were removed from half of the plants. Plants whose mature pods were removed produced significantly greater numbers of flowers, ovules and viable seeds and a greater total weight of pods than did untreated plants. However, the proportion of ovules giving rise to viable seeds and to aborted seeds and undeveloped ovules was the same in plants of both groups. Approximately 50% of ovules of marketable pods, in all plants, yielded viable seeds. The 12 cohorts of flowers contributed markedly different amounts to plant yield. Early and later cohorts contributed much less than did middle cohorts in terms of marketable pod dry weight and numbers of fully developed seeds, aborted seeds, and undeveloped ovules. Peak productivity was therefore obtained from the middle phase of flowering in these annual bean plants. The lower pod yield of later cohorts is interpreted as a result of competition for limited resources between maturing pods and new flowers. We consider possible causes for the various fates of flowers and ovules and discuss flowers as plant modules suitable for demographic examination.     

Resource patch size and flexible foraging in white-faced capuchins (Cebus capucinus)

White-faced capuchins (Cebus capucinus)on Barro Colorado Island, Panama, have a flexible foraging strategy. Typically, foraging party size is small and individuals feed dispersed from one another. When seasonal fruiting of large volume trees occurs, the majority of the group forages simultaneously. As C. capucinusdo not display a rigorous dominance structure and there are few indications that individuals or coalitions monopolize food patches,individuals are expected to display scramble strategies instead of high frequencies of contest competition. I recorded foraging party size (simultaneous foragers), the total number of animals to feed successively, and the diameter at breast height (DBH) of fruit trees used in two habituated troops. Individuals in each group spent a substantial amount of time — 65 and 48% of foraging time for each group — foraging in party sizes of one. Monkeys predominantly foraged alone in small trees (0- to 20- cm DBH), successively in medium trees (21- to 60- cm DBH), and simultaneously in large trees (>61- cm DBH). They used small trees more frequently than all other tree classes. In medium-sized trees, although fruit was plentiful, space was limited. In these trees Cebusforaged successively. In large-volume trees, space and fruit were abundant and several individuals fed together. As the DBH of fruiting trees increased, the average foraging party size increased exponentially. Cebus capucinusat Barro Colorado Island modify their foraging party size to adapt to the seasonal patterns of fruit production.     

Does the morphology of animal foraging pits influence secondary seed dispersal by ants?

Secondary seed dispersal by ants (myrmecochory) is an important process in semi‐arid environments where seeds are transported from the soil surface to an ant nest. Microsites from which ants often remove seeds are the small pits and depressions made by native and exotic animals that forage in the soil. Previous studies have demonstrated greater seed retention in the pits of native than exotic animals, but little is known about how biotic factors such as secondary seed dispersal by ants affect seed removal and therefore retention in these foraging pits. We used an experimental approach to examine how the morphology of burrowing bettong (Bettongia lesueur), greater bilby (Macrotis lagotis), short‐beaked echidna (Tachyglossus aculeatus) and European rabbit (Oryctolagus cuniculus) foraging pits and ant body size influenced ant locomotion and seed removal from pits along an aridity gradient. Ants took 3.7‐times longer to emerge from echidna pits (19.6 s) and six‐times longer to emerge from bettong pits (30.5 s) than from rabbit pits (5.2 s), resulting in lower seed removal from bettong pits than other pit types. Fewer seeds were removed from pits when cages were used to exclude large body‐sized (>2 mm) ants. Few seeds were removed from the pits or surface up to aridity values of 0.5 (humid and dry sub‐humid), but removal increased rapidly in semi‐arid and arid zones. Our study demonstrates that mammal foraging pit morphology significantly affects ant locomotion, the ability of ants to retrieve seeds, and therefore the likelihood that seeds will be retained within foraging pits.     

Assessing the effects of elephant foraging on the structure and diversity of an Afrotropical forest

African forest elephants (Loxodonta cyclotis) are ecosystem engineers that browse and damage large quantities of vegetation during their foraging and movement. Though elephant trail networks and clearings are conspicuous features of many African forests, the consequences of elephant foraging for forest structure and diversity are poorly documented. In this study in northeastern Gabon, we compare stem size, stem density, proportional damage, species diversity, and species relative abundance of seedlings and saplings in the vicinity of seven tree species that produce elephant-preferred fruits (“elephant trees”) relative to control trees that do not. Across 34 survey trees, with a combined census area of 2.04 ha, we recorded data on 26,128 woody stems in three sizes classes. Compared with control trees, the area around elephant trees had the following: (a) a significantly greater proportion of damaged seedlings and a marginally greater proportion of damaged saplings (with 82% and 24% greater odds of damage, respectively); (b) no significant difference in stem density or species diversity; and (c) a significantly greater relative abundance of seedlings of elephant tree species. Increasing distance away from focal elephant trees was associated with significantly reduced sapling stem damage, significantly increased sapling stem density, and significantly increased sapling species diversity. Considered in sum, our results suggest that elephants can affect the structure and diversity of Afrotropical forests through their foraging activities, with some variation based on location and plant size class. Developing a more complete understanding of elephants’ ecological effects will require continued research, ideally with manipulative experiments.     

A mechanism for resource allocation among sympatric heteromyid rodent species

Summary Laboratory feeding experiments were conducted with Dipodomys ordii and Perognathus flavus in an attempt to discover a mechanism which might result in seed size selection. There was no marked difference in the proportions of four seed types collected whether the rodents foraged in the presence or absence of one another. However, analysis of the variability in weight of each of the seed types collected by the two species showed that when alone, the larger kangaroo rat was less effective at harvesting all of a uniformly distributed mixture of seeds. When in the presence of one another both species could harvest enough of the mixed, uniformly distributed seed to coexist indefinitely, but when the food source was presented as four large clumps the kangaroo rat's foraging effectiveness increased tremendously so that the pocked mouse was almost entirely unable to harvest any seed. These data, in light of mobility differences between large and small heteromyids, suggest a mechanism whereby the larger, more mobile kangaroo rats forage for the most readily available (large or clumped) seeds over a relatively large area. The smaller pocket mice, by virtue of their relative efficiency in harvesting seeds can utilize the less detectable seeds which are energetically too demanding for the larger kangaroo rats to harvest. Behavioral dominance of the larger animals may help prevent the smaller from utilizing the most readily available seeds. The patterns of seed size and foraging site selection described in the literature may be easily accounted for by this difference in foraging strategy.