Ecological niche differences betweenAlouatta palliata andCebus capucinus comparing feeding modes,branch use,and diet

During the wet season, two sympatric species of primates,Alouatta palliata (mantled howlers) andCebus capucinus (white-faced capuchins), were assayed for feeding niche differences through behavioral and habitat use patterns at Refugio de Fauna Silvestre Curu in Costa Rica. Differences in the use of relative diameter and thickness of branches and five different modes of feeding were compared between the species. White-faced capuchins used more manipulative modes of obtaining food, a wider range of arboreal habitat, and had a more diverse diet than mantled howlers. Mantled howlers may be more restricted than white-faced capuchins in arboreal microhabitat use due to their possible need for large support branches during feeding bouts and resting periods. We report that differences in feeding behaviors, diet, and arboreal habitat use seem to play a large role in separating these species niches.     

High density of white-faced capuchins (Cebus capucinus) and habitat quality in the Taboga Forest of Costa Rica

Across the globe, primates are threatened by human activities. This is especially true for species found in tropical dry forests, which remain largely unprotected. Our ability to predict primate abundance in the face of human activity depends on different species' sensitivities as well as on the characteristics of the forest itself. We studied plant and primate distribution and abundance in the Taboga Forest, a 516-ha tropical dry forest surrounded by agricultural fields in northwestern Costa Rica. We found that the density of white-faced capuchins (Cebus capucinus) at Taboga is 2–6 times higher than reported for other long-term white-faced capuchin sites. Using plant transects, we also found relatively high species richness, diversity, and equitability compared with other tropical dry forests. Edge transects (i.e., within 100 m from the forest boundary) differed from interior transects in two ways: (a) tree species associated with dry forest succession were well-established in the edge and (b) canopy cover in the edge was maintained year-round, while the interior forest was deciduous. Sighting rates for capuchins were higher near water sources but did not vary between the edge and interior forest. For comparison, we also found the same to be true for the only other primate in the Taboga Forest, mantled howler monkeys (Alouatta palliata). Year-round access to water might explain why some primate species can flourish even alongside anthropogenic disturbance. Forest fragments like Taboga may support high densities of some species because they provide a mosaic of habitats and key resources that buffer adverse ecological conditions.     

Down by the riverside: Riparian edge effects on three monkey species in a fragmented Costa Rican forest

Rivers represent natural edges in forests, serving as transition zones between landscapes. Natural edge effects are important to study to understand how intrinsic habitat variations affect wildlife as well as the impact of human-induced forest fragmentation. We examined the influence of riparian and anthropogenic edge on mantled howler, white-faced capuchin, Central American spider monkeys, and vegetation structure at La Suerte Biological Research Station (abbreviated as LSBRS), Costa Rica. We predicted lower monkey encounter rate, tree species richness, and median dbh at both edge types compared to interior and that monkeys would show species-specific responses to edge based on size and diet. We expected large, folivorous–frugivorous howler monkeys and small, generalist capuchins would be found at increased density in forest edge, while large, frugivorous spider monkeys would be found at decreased density in forest edge. We conducted population and vegetation surveys along interior, riparian, and anthropogenic edge transects at LSBRS and used GLMM to compare vegetation and monkey encounter rate. Tree species richness and median dbh were higher in forest interior than anthropogenic edge zones. Although spider monkey encounter rate did not vary between forest edges and interior, howler monkeys were encountered at highest density in riparian edge, while capuchins were encountered at highest density in anthropogenic edge. Our results indicate that diverse forest edges have varying effects on biota. Vegetation was negatively affected by forest edges, while monkey species showed species-specific edge preferences. Our findings suggest that riparian zones should be prioritized for conservation in Neotropical forests.     

Predictors of Parasitism in Wild White-Faced Capuchins (Cebus capucinus)

Parasite infections in wildlife are influenced by many factors including host demography, behavior and physiology, climate, habitat characteristics, and parasite biology and ecology. White-faced capuchins (Cebus capucinus) host a suite of gastrointestinal and pulmonary parasites, yet the mechanisms affecting host susceptibility and parasite transmissibility have not been examined in this host species. We used the information-theoretic approach (Akaike’s information criterion) and traditional null-hypothesis testing to determine which host characteristics, behaviors, or environmental factors affected the presence of two prevalent capuchin parasites (Filariopsis barretoi and Strongyloides sp.) as well as parasite species richness in four groups of wild capuchins from September 2007 to January 2008 and January to August 2009. Older capuchins were more likely to be infected with Filariopsis barretoi and had higher parasite species richness. Age-biased nematode infections may stem from age differences in capuchin behavior and physiology while high species richness likely results from long- term exposure to numerous parasite species. Infections with Strongyloides sp. were more likely to occur in the dry season when capuchins often descend to the forest floor to drink from terrestrial water sources, potentially increasing their risk of infection from soil-borne larvae. Capuchin behaviors were poor predictors of parasitism, as were female rank, host sex, home range size, and habitat quality. Many of our results were atypical for primate parasitology, suggesting that host–parasite interactions, and subsequently infection risk, may differ in highly seasonal habitats such as tropical dry forests where these monkeys occur.     

The influence of anthropogenic edge effects on primate populations and their habitat in a fragmented rainforest in Costa Rica

When a forest is fragmented, this increases the amount of forest edge relative to the interior. Edge effects can lead to loss of animal and plant species and decreased plant biomass near forest edges. We examined the influence of an anthropogenic forest edge comprising cattle pasture, coconut plantations, and human settlement on the mantled howler (Alouatta palliata), white-faced capuchin (Cebus capucinus), Central American spider monkey (Ateles geoffroyi), and plant populations at La Suerte Biological Research Station (LSBRS), Costa Rica. We predicted that there would be lower monkey encounter rate, mean tree species richness, and diameter at breast height (DBH) in forest edge versus interior, and that monkeys would show species-specific responses to edge based on diet, body size, and canopy height preferences. Specifically, we predicted that howler monkeys would show positive or neutral edge effects due to their flexible folivorous diet, large body size, and preference for high canopy, capuchins would show positive edge effects due to their diverse diet, small body size, and preference for low to middle canopy, and spider monkeys would show negative edge effects due their reliance on ripe fruit, large body size, and preference for high upper canopy. We conducted population and vegetation surveys along edge and interior transects at LSBRS. Contrary to predictions, total monkey encounter rate did not vary between the forest edge and forest interior. Furthermore, all three species showed neutral edge effects with no significant differences in encounter rate between forest edge and interior. Interior transects had significantly higher mean tree species richness than edge transects, and interior trees had greater DBH than edge trees, although this difference was not significant. These results suggest that forest edges negatively impact plant populations at La Suerte but that the monkeys are able to withstand these differences in vegetation.     

Stable isotope ratios indicate diet and habitat use in New World monkeys

This paper demonstrates the use of stable isotope ratios of carbon and nitrogen in animal tissues for indicating aspects of species behavioral strategy. We analyzed hair from individuals representing four species of New World monkeys (Alouatta palliata, the mantled howler; Ateles geoffroyi, the spider monkey; Cebus capucinus, the capuchin; and Brachyteles arachnoides, the woolly-spider monkey or muriqui) for δ¹³C and δ¹⁵N using previously developed methods. There are no significant differences in either carbon or nitrogen ratios between sexes, sampling year, or year of analysis. Seasonal differences in δ¹³C reached a low level of significance but do not affect general patterns. Variation within species was similar to that recorded previously within single individuals. The δ¹³C data show a bimodal distribution with significant difference between the means. The two monkey populations living in an evergreen forest were similar to each other and different from the other two monkey populations that inhabited dry, deciduous forests. This bimodal distribution is independent of any particular species' diet and reflects the level of leaf cover in the two types of forest. The δ¹⁵N data display three significantly different modes. The omnivorous capuchins were most positive reflecting a trophic level offset. The spider monkeys and the muriquis were similar to one another and significantly more positive than the howlers. This distribution among totally herbivorous species correlates with the ingestion of legumes by the howler monkey population. In combination, these data indicate that museum-curated primate material can be analyzed to yield information on forest cover and diet in populations and species lacking behavioral data. Am. J. Phys. Anthropol. 103:69–83, 1997. © 1997 Wiley-Liss, Inc.     

The effects of provisioning and crop-raiding on the diet and foraging activities of human-commensal white-faced capuchins (Cebus capucinus)

Non-human primates are coming into increasingly frequent contact with humans and with human-modified environments. The potential for monkeys to survive in such modified landscapes is questionable, and is likely related to a species' behavioral plasticity, particularly as it relates to diet. In this study, I explore the ways in which white-faced capuchins (Cebus capucinus) adjust their diet and foraging behaviors in response to anthropogenic impact. I compare a troop of human-commensal monkeys and a similar troop of wild-feeding monkeys living within the Curú Wildlife Refuge in western Costa Rica for differences in overall diet composition and activity budgets to evaluate the impact of habitat change in this context. The commensal-living white-faced capuchins rely on raided coconut (Cocos nucifera) and oil palm (Elaeis guineensis) crops and provisioned or stolen human foods for over one-half of their total diet. Regardless of this highly anthropogenic diet, the two study troops do not significantly differ in their activity budgets, and the human-commensal troop maintains wild-foraging activities consistent with those of the wild-feeding troop. These data suggest that the white-faced capuchins at this site are responding to anthropogenic disturbance primarily through the exploitation of human food resources, but they do not yet appear to have lost the foraging skills required to survive in this modified landscape on their own. This study adds to our growing body of knowledge on primate survival in matrix habitats, and will hopefully inform primate management plans throughout the Neotropics.     

Crop Feeding by Brown Howlers (<Emphasis Type="Italic">Alouatta guariba clamitans</Emphasis>) in Forest Fragments: The Conservation Value of Cultivated Species

Primates inhabiting human-modified habitats often complement their diets with cultivated species. Although this flexible foraging behavior reduces feeding stress in animals inhabiting small or low-quality habitats, its potential economic costs may promote negative human–nonhuman primate interactions. It is critical to assess the importance of cultivated species to primate diets, the factors influencing their exploitation, and their associated economic costs to evaluate the conservation value of cultivated species and the impact of crop feeding on human–nonhuman primate coexistence. Based on a 30-months study of three groups of brown howlers (Alouatta guariba clamitans) inhabiting <10-ha forest fragments in Rio Grande do Sul State, southern Brazil, and semistructured interviews with landowners, we assessed 1) the cultivated species used as food sources, 2) the relationship between food availability and crop feeding, and 3) the potential economic costs of crop feeding and their impact on landowner–howler coexistence. Brown howlers exploited six cultivated species as fruit or seed sources. This exploitation was concentrated on ripe fruit of Psidium guajava, Citrus reticulata, Diospyros kaki, and Eriobotrya japonica, which together accounted for up to 32% of total feeding records during some months. Availability predicted fruit consumption for three of these species. The overall availability of cultivated fruit also predicted its consumption, whereas overall wild fruit availability did not. Despite potential economic costs of crop feeding (US$320–US$1141 per year), landowners reported no conflict with howlers arising from their behavior, an amicable response that supports the conservation value of cultivated plants in different landscape elements of the anthropogenic matrix.     

Social Aspects of Fur-rubbing in <Emphasis Type="Italic">Cebus capucinus</Emphasis> and <Emphasis Type="Italic">C. apella</Emphasis>

Capuchins rub particular plant materials into their pelage, a behavior for which most authors have proposed a medicinal function (Baker in American Journal of Primatology 38:263–270, 1996, Baker, M. (1998). Fur Rubbing as Evidence for Medicinal Plant Use by Capuchin Monkeys (Cebus capucinus): Ecological, Social, and Cognitive Aspects of the Behavior. Dissertation thesis. University of California Riverside; DeJoseph et al. in Journal of the American Academy of Dermatology 46:924–925, 2002). Individuals fur-rub solitarily or in groups, but researchers have not conducted studies to understand the differences. We investigated the link between the form of fur-rubbing and other social variables in 2 capuchin species. We supplied 2 captive groups —white-faced capuchins and tufted capuchins— with citrus fruit and onions and explored the behavioral processes and social aspects underlying the activity. We documented the occurrence, number of participants, and effect of recruitment behaviors in fur-rubbing subgroups. We investigated the role of kinship, affiliation, and dominance relationships in accounting for fur-rubbing groups. There is a significant difference in the form of fur-rubbing between white-faced and tufted capuchins. White-faced capuchins fur-rubbed mainly in subgroups and performed a particular behavior to recruit prospective participants, whereas tufted capuchins fur-rubbed mainly alone, and showed no particular motivation to be joined by other group members. White-faced capuchins could fur-rub together frequently, whatever their degree of kinship, affiliation, or dominance interval. In tufted capuchins, fur-rubbing appeared to be significantly affected by kinship and dominance.     

Compound grips in tufted capuchin monkeys (Sapajus spp and Sapajus libidinosus)

An experimental study with captive individuals and study of video recordings of wild monkeys explored whether and how tufted capuchin monkeys use onehand to hold one or more objects with multiple grips (compound grips). A task designed to elicit compound grip was presented to five captive tufted capuchin monkeys (Sapajus spp). The monkeys held one to four balls in onehand and dropped the balls individually into a vertical tube. Multiple simple grips and independent digit movements enabled separate control of multiple objects in one hand. Monkeys always supported the wrist on the horizontal edge of the tube before releasing the ball. Increasing the number of balls decreased the likelihood that the monkeys managed the task. Wild bearded capuchins (Sapajus libidinosus) used compound grips spontaneously to store multiple food items. Compound grips have been described in macaques, gorillas, chimpanzees, and humans, and now in a New World primate. We predict that any primate species that exhibits precision grips and independent digit movement can perform compound grips. Our findings suggest many aspects of compound grip that await investigation.     

Olfactory discrimination of urine odors from five species by tufted capuchin (Cebus apella)

Urine collected from New World monkeys (tufted capuchin, squirrel monkey, cotton-top tamarin) and Old World monkeys (rhesus macaque, Japanese macaque), was used as the odor stimuli. Two adult tufted capuchins were trained on a successive odor-discrimination task with two odors, 30 trials each, in one session per day. Responses to one of the two odors (S+) were reinforced by sweet water. The monkeys failed to discriminate between the urine from the two species of macaques but could discriminate among the urine from the three species of New World monkeys. Furthermore, similarity of urine odors was analyzed by multi-dimensional scaling (MDS) and a cluster analysis. These analysis suggested that the tufted capuchin can distinguish differences among New World monkeys but not between the macaques. The natural distribution of the tufted capuchin overlaps with that of other New World monkeys, but it does not overlap with those of Old World monkeys. Consequently, it can be concluded that this difference in olfactory recognition in the tufted capuchin reflects their sympatric and allopatric relationships with other species.     

Absence of Howlers (<Emphasis Type="Italic">Alouatta palliata</Emphasis>) Influences Tree Seedling Densities in Tropical Rain Forest Fragments in Southern Mexico

The disappearance of frugivorous primates in fragmented forests can potentially lower the rates of seed dispersal and recruitment of endozoochorous tree species, thus altering plant community structure. We quantified seedling density for 7 tree species that are common in the feces of mantled howlers (Alouatta palliata) in 6 rain forest fragments in northern Chiapas, Mexico. Howlers were present in 3 of the fragments and absent in the other 3. We compared seedling density in primate sleeping sites in inhabited fragments with control sites, which were structurally similar to sleeping sites but where we did not find monkey feces, in both inhabited and uninhabited fragments. For each tree species, we determined the relationship between seedling density and the local density of seeds and adult trees. In fragments where howlers were present, seedling density for 4 of the focal tree species (Brosimum alicastrum, Dialium guianense, Manilkara zapota, and Nectandra ambigens) was greater in sleeping sites than in control sites found in the same fragments. Moreover, seedling density of Dialium guianense was greater in the control sites of fragments inhabited by howlers than in fragments where this primate is absent. Seedling density of these 4 species correlates positively with seed density on the forest floor; however, we observed no correlations between seedling density and the density of adult trees. Our results suggest that the diversity of the seedling community of tree species dispersed by howlers may decline in fragments where this seed disperser is absent. These findings, together with the fact that only 5% of the study region is currently covered by forest and 81% of the forest remnants are uninhabited by mantled howlers, suggest that the potential long-term recovery of associated populations of tropical tree species dispersed by this primate species is highly uncertain. Conservation and restoration efforts should be aimed at restoring or replacing the ecological role played by this important seed disperser in the region.     

Picture processing in tufted capuchin monkeys (Cebus apella)

Although pictures are frequently used in place of real objects to investigate various aspects of cognition in different non-human species, there is little evidence that animals treat pictorial stimuli as representations of the real objects. In the present study, we carried out four experiments designed to assess picture processing in tufted capuchin monkeys (Cebus apella), using a simultaneous Matching-to-Sample (MTS) task. The results of the first three experiments indicate that capuchins are able to match objects with their colour photographs and vice versa, and that object-picture matching in this New World monkey species is not due to picture-object confusion. The results of the fourth experiment show that capuchins are able to recognize objects in their pictures with a high level of accuracy even when less realistic images, such as black-and-white photographs, silhouettes and line drawings, are employed as bi-dimensional stimuli. Overall, these findings indicate that capuchin monkeys are able to establish a correspondence between the real objects and their pictorial representations.     

Neotropical Primates in a Regenerating Costa Rican Dry Forest: A Comparison of Howler and Capuchin Population Patterns

Few data exist on how primate populations return to regenerating tropical forests. We compare the ways that two populations of neotropical monkeys, Alouatta palliata and Cebus capucinus, expanded over a 28-year period after the establishment of Santa Rosa National Park on reclaimed ranchlands in Costa Rica. We found that both howler and capuchin populations increased substantially in size subsequent to protection, but the howler population grew faster. This is likely due to their faster-paced life-history pattern than that of capuchins. The howler population increased mainly via the establishment of many new groups, whereas the capuchins expanded mainly by increasing the size of existing groups. We related this finding to the fact that capuchins are limited largely by their need to drink from water holes during the dry seasons whereas howlers are limited principally by their preference for larger-sized trees that occur in older forests. Proportions of adult male capuchins increased significantly during our study, likely due to skewed sex ratio at birth or male-biased immigration into the protected park or both factors. Our main finding is that, in as short a time period as 28 years, we can substantially enhance the size of monkey populations by allowing the regeneration of tropical forest. Furthermore, we provide a preliminary interpretation of how extrinsic factors—deforestation, hunting, crop-spraying, destruction of the watershed—and intrinsic variables, e.g., pace of reproduction; diet, differentially affect not only each species' vulnerability to extinction but also its capacity to recover when human disturbances are minimized.     

Behavioral indicators of ovarian phase in white-faced capuchins (Cebus capucinus)

In many primate species, conspicuous behavioral and/or morphological changes are indicators of the fertile phase of the female cycle. However, several primate species, such as the white-faced capuchin, lack these cues. This is referred to as "concealed ovulation," and is argued to be a reproductive strategy that confuses paternity and lowers the risk of infanticide. We studied 10 adult female white-faced capuchins in Santa Rosa National Park, Costa Rica, from January to June 2002. We determined their ovarian cycling patterns by analyzing fecal ovarian hormones, and compared simultaneously collected behavioral data to determine which, if any, cues females use to signal their fertile phases. We found that four females cycled during the study period but ceased to cycle without becoming pregnant. We considered several explanations for the lack of conception during our study, including reproductive seasonality. We found that female C. capucinus showed only small increases in rates of affiliative/proceptive behaviors directed toward adult males during their periovulatory phases. The best indicator of cycle phase was a significant increase in male affiliative behaviors (e.g., following and grooming bouts) and sexual behaviors (e.g., copulations and courtship displays) directed toward females during the periovulatory phase compared to the nonovulatory phase. Our finding that females exhibit little proceptive behavior, but that copulations and male courtship are nonetheless concentrated in periovulatory phases suggests that even though females do not provide behavioral and morphological cues to ovulation, males are still able to detect it. Infanticide occurs with some frequency in these monkeys, and there is evidence for postconceptive mating as a female strategy to lower risk of infanticide via paternity confusion. However, despite this occurrence of nonconceptive mating and the absence of female cues to ovulation, truly concealed ovulation does not appear to be characteristic of this study population of white-faced capuchins.     

The Repertoire and Social Function of Facial Displays in Cebus capucinus

Systematic studies on facial displays in capuchins are limited and based mainly on studies of tufted capuchins (Cebus apella). Despite the great social-morphological variability within Cebus suggesting possible morphological and functional variations in the facial displays of different species, no study has considered thoroughly visual communication in the genus. Our aim was to describe the facial displays of white-faced capuchins and to assess their distribution and communicative function. We observed 15 captive white-faced capuchins in the Primate Centre of the Louis Pasteur University of Strasbourg, for a total of 198 h. We described the following facial displays: relaxed open-mouth, lip-smacking, open-mouth threat-face, silent bared-teeth, open-mouth silent bared-teeth, protruded-lip face, and tongue-out. We never observed the scalp-lifting display, one of the most common displays characterizing tufted capuchins. White-faced capuchins use the majority of facial displays in an affiliative or playful context; only the open-mouth threat-face display is associated with aggressive behaviors. White-faced capuchins lack ritualized signals of submission. The fact that in white-faced capuchins the silent bared-teeth display conveys only a positive message, while in tufted capuchins it signals submission as well as affiliation, supports the covariation hypothesis (Thierry 2004 Social epigenesis. In B. Thierry, M. Singh, & W. Kaummans (Eds.), Macaque societies: A Model for the study of social organization, pp. 267–294. Oxford University Press).     

Endowment effect in capuchin monkeys

In humans, the capacity for economically rational choice is constrained by a variety of preference biases: humans evaluate gambles relative to arbitrary reference points; weigh losses heavier than equally sized gains; and demand a higher price for owned goods than for equally preferred goods that are not yet owned. To date, however, fewer studies have examined the origins of these biases. Here, we review previous work demonstrating that human economic biases such as loss aversion and reference dependence are shared with an ancestrally related New World primate, the capuchin monkey (Cebus apella). We then examine whether capuchins display an endowment effect in a token-trading task. We identified pairs of treats (fruit discs versus cereal chunks) that were equally preferred by each monkey. When given a chance to trade away their owned fruit discs to obtain the equally valued cereal chunks (or vice versa), however, monkeys required a far greater compensation than the equally preferred treat. We show that these effects are not due to transaction costs or timing issues. These data suggest that biased preferences rely on cognitive systems that are more evolutionarily ancient than previously thought-and that common evolutionary ancestry shared by humans and capuchins may account for the occurrence of the endowment effect in both species.     

Carrying and hand-use patterns in Panamanian white-faced capuchins (Cebus capucinus)

The aim of this project was to examine the potential influence of postural regulation on capuchin hand-use patterns by focusing on tasks that involved the carrying of objects. Two months were spent on Barro Colorado Island, Panama, collecting data on ten white-faced capuchins (Cebus capucicus) (four adult males/six adult females). Over 215 contact hours were spent with the monkeys, and a total of 213 carries were recorded. No hand preference bias at the population level was found for the carry task; and no detectable hand-use patterns for carry were correlated with the weight of the object carried (except a right-hand bias for carrying medium-sized objects), the plane of the monkey's movement, its locomotor pattern, or its height above the ground. Therefore, the results of this study do not support the hypothesis that the need for postural regulation while carrying an object influences hand-use patterns in free-ranging capuchins. Our results are especially important because they are contrary to the results of the only other free-ranging capuchin laterality study conducted to date (Panger, 1998).     

Extraction of hermit crabs from their shells by white-faced capuchin monkeys (<Emphasis Type="Italic">Cebus capucinus</Emphasis>)

We observed two capuchin monkeys (Cebus capucinus) feeding on hermit crabs (Coenobita compressus) on the coast, and the tactics they used to extract this well-protected prey. The observations took place during the dry season at Playa Escondida beach, Puntarenas, Costa Rica. The capuchins descended from trees at the back edge of the beach to capture passing hermit crabs. Both capuchins extracted the hermit crabs from their protective shells by holding the shell with one hand and pulling the crab out with the other. Even though this was accomplished within seconds, the extraction of hermit crabs from their shells did not appear to be a straightforward task. Once the capuchins succeeded in pulling the crabs out of their shells, they consumed the soft abdomen and discarded the rest of the crab's body. To our knowledge, the consumption of hermit crabs has not been previously reported for any capuchin monkey (Cebus or Sapajus). Our observations provide a new example of extractive foraging by capuchins, and thus an additional natural context for which fine motor skills (which are highly developed in capuchins) are necessary.     

Tool use in Cebus

This paper summarizes early anecdotal information and systematic studies of tool use in capuchin monkeys (Cebus spp.). Tool use in capuchins is neither context specific nor stereotyped. The success of capuchins in using tools and in exploiting a variety of food resources in the wild derives from several factors: their manipulative abilities, interest in external objects and a tendency to explore the environment. In using tools, capuchins are similar to apes and more proficient than other monkey species. A cognitive approach indicates, however, that (in contrast with chimpanzees) they never develop an understanding of the requirements of the tool tasks presented.