Ontogeny of provisioning in two populations of wild golden lion tamarins (Leontopithecus rosalia)

Provisioning weaned young is an important part of the cooperative rearing system of marmosets and tamarins (family: Callitrichidae). Juvenile callitrichids receive a substantial proportion of their diet from all adult group members, whereas juveniles of most other primate species only receive food from the mother infrequently via scrounging. We conducted a longitudinal study of provisioning to 21 young wild golden lion tamarins through their first year of life in two Brazilian reserves. Juveniles were predicted to experience higher provisioning rates at the site with mature forest (the presumed ancestral habitat) and in territories that contained a higher proportion of preferred habitat within the mature forest reserve. Key measures of provisioning did not differ substantially between habitat types. The rates at which juveniles begged for food were very similar across habitats, while the influence of habitat on the rates at which juveniles received food was small and varied across age groups. The most pronounced differences between the reserves were: adult resistance to food-transfer attempts was less frequent, food-offering calls before food transfer were more frequent, and the proportion of prey provisioned was higher in the reserve in which earlier successional-stage forest predominated. Within the mature forest reserve, begging success of young juveniles covaried positively with preferred habitat. Rather than overt caretaker–offspring conflict, an information–feedback loop between juveniles and adults appears to drive provisioning changes as juveniles mature. Direct measures of resource abundance are needed to help clarify the mechanisms by which forest successional stage influences provisioning.     

Costs and benefits of allomothering in wild capped langurs (Presbytis pileata)

Summary Data from a 15-month field study of the capped langur monkey, Presbytis pileata, in Bangladesh indicate that allomothering behavior is restricted to particular female-infant dyads. Primary allomothers were all parous adult females; nulliparous females rarely allo-mothered, Newborn infants were transferred from the mother to other females within a short time of birth, as reported for some other colobine species, but over the first 3 months of life each neonate's contact with nonmothers was largely restricted to a single allomother in each of five study groups. Capped langur mothers with newborn infants spent more time feeding when the infant was being allomothered than when it was in the mother's care. The model of allomothering as a selfish behavior by nulliparous or pregnant females used to enhance maternal skills at the expense of mothers is not supported by this study. Rather, allomothering may have adaptive significance as altruistic behavior among group females, in that it enables lactating females to increase feeding time. Capped langur allomothering is best interpreted as a low-cost behavior that can benefit recipients that may or may not be related. Offprint requests to the present address     

Costs and benefits of capsaicin-mediated control of gut retention in dispersers of wild chilies

A fundamental way in which animal-dispersed plants can influence the viability and distribution of dispersed seeds is through control of retention time in the guts of dispersers. Using two species of wild chilies and their dispersers, we examined how chemical and physical properties of fruits and seeds mediate this interaction. Capsicum chacoense is polymorphic for pungency, occurs in Bolivia, and is dispersed mostly by elaenias. Capsicum annuum is not polymorphic, occurs in Arizona (USA), and is dispersed mostly by thrashers. We first tested whether capsaicin, the substance responsible for the pungency of chilies, affects gut retention time of seeds in primary dispersers. Capsaicin slowed gut passage of seeds but did so in a manner that differed greatly between bird species because the constipative effects of capsaicin occurred only after an 80-minute time lag. Elaenias in Bolivia held only 6% of C. chacoense seeds for > 80 minutes, whereas thrashers in Arizona held 78% of C. annuum seeds for > 80 minutes. Next we examined the effects of retention time on seed viability and germination. Increased retention resulted in a greater proportion of seeds germinating in C. annuum, had no effects on non-pungent C. chacoense, and had negative effects on pungent C. chacoense. These divergent effects are explained by differences in seed coat morphology: seed coats of pungent C. chacoense are 10-12% thinner than those of the other two types of seeds. Thus, longer retention times damaged seeds with the thinnest seed coats. In C. annuum, seed viability remained high regardless of retention time, but germination increased with retention, suggesting a role for scarification. Thus, in C. annuum, fruit chemistry appears well matched with seed morphology and disperser physiology: capsaicin extends gut retention for most seeds, resulting in greater seed scarification and higher germination rates. Increased retention of pungent C. chacoense seeds is detrimental, but because the primary consumers have short retention times, capsaicin slows only a small proportion of seeds, minimizing negative effects. These results illustrate the importance of context in studies of fruit secondary metabolites. The same chemical can have different impacts on plant fitness depending on its morphological, physiological, and ecological context.     

Prey-capture benefits in a mixed-species group of Amazonian tamarins,Saguinus fuscicollis and S. mystax

Summary Tamarins of the genus Saguinus feed on a wide range of arthropods and small vertebrates, which compose a critical component of their diet. This paper examines the foraging patterns and capture success of the Avila-Peres saddle-back (S. fuscicollis avilapiresi) and the red-capped moustached tamarin (S. mystax pileatus) in very stable mixed-species groups, and whether and how any foraging benefits for either species resulted from their association. Moustached tamarins actively searched for prey items which were mainly well exposed on the midstorey foliage. Saddle-back tamarins, on the other hand, foraged at lower heights, largely by manipulating a variety of microhabitats potentially concealing embedded prey. The foraging activity of the numerically dominant and larger-bodied moustached tamarins often resulted in prey items escaping to lower substrates, usually the forest leaf-litter. The beating effect of this species substantially facilitated captures of large, mobile prey items by saddle-backs, which were highly adept at locating and retrieving flushed prey. It is estimated that, while saddle-backs obtained 66–73% of their prey biomass from flushed items, this proportion was substantially lower (2–9%) for moustached tamarins. Commensal insectivory appears to involve a highly asymmetric benefit to saddle-backs, and a low cost to moustached tamarins, which partly explains the stability of mixed-species groups. Correspondence to the present address     

Costs and benefits of brood desertion in female kentish plovers,Charadrius alexandrinus

Female kentish plovers Charadrius alexandrinus typically desert their broods after the chicks hatch, i.e. 1–4 weeks before the chicks fledge or become independent. In this paper we investigate the costs and benefits of desertion for females. Desertion incurs a cost for females: following desertion chick survival in broods is lower (0.95 +- 0.02 day^–1) than before the female deserts (0.98 +- 0.01 day^–1). We investigated several possible causes for reduced brood survival by comparing characteristics of broods before and after desertion (controlling for differences in brood age). After desertion males increased the time they spent foraging and they tended to reduce time spent brooding chicks. Increased mortality of chicks may occur in deserted broods because following desertion (1) males spend less time alert in vigilance behaviour than before desertion, (2) they attend the chicks from greater distances, and (3) they show greater distraction display distances (in response to human intruders). Growth or development of chicks, measured by weight gain and tarsus length, was not different before and after desertion. Females gain two potential benefits from desertion: (1) they may remate and produce a second brood within the same breeding season or (2) they may enhance their probability of surviving to breed in a subsequent season. At least 27% of female kentish plovers that deserted remated and renested in the same season in this study. In contrast, we found no evidence that brood desertion increased the survival of females: there was no difference in local survival rate (return rate) for females deserting before or after 6 days brood age. These results clearly demonstrate that female kentish plovers that desert their offspring prior to fledging incur costs, but we suggest that there is a trade-off with the potential benefits gained by remating and making a second breeding attempt in the same season.     

Costs, benefits, and plasticity of construction of nest defensive structures in paper wasps

Various animals build nests with defensive structures to deter predation on offspring. Construction of nest defensive structures can reduce the probability of predation but will involve various costs. Here, we examined both the costs and benefits of the construction of a nest defensive structure in a paper wasp, Polistes chinensis antennalis, and clarify whether the paper wasp changes the level of defensive structures of nests depending on predation risk. A foundress (queen) of the paper wasp starts a colony in spring and maintains her nest alone until the emergence of workers. At this stage, pupae in the nests are sometimes preyed on by conspecifics of other nests. The intruder needs to break the cocoon, which seals the entrance of the cell, to extract the pupa from the cell. Foundresses often apply nest material (pulp) to the surface of cocoons in their nests. We found that pulp on a cocoon increased the time an intruder required to break the cocoon. This result shows that the pulp structure on cocoons helps to prevent predation on pupae. On the other hand, pulp on cocoons involved costs, including time required to collect pulp and being a potential obstacle to the emergence of workers from the cocoon. Additionally, we found that the amount of pulp on cocoons was greater in nests under higher predation risk than nests under lower predation risk. These results suggest that pulp on cocoons is a nest defensive structure, and foundresses adjusted the construction of the defensive structure depending on predation risk.     

Facultative polyandry and the role of infant-carrying in wild saddle-back tamarins (Saguinus fuscicollis)

Summary Wild saddle-back tamarins (Saguinus fuscicollis) in southeastern Peru have a variable mating system that can differ both between territories at any one time and within territories over time. Groups are usually monogamous or cooperatively polyandrous, but are occasionally even polygynous. This study addressed the following questions: why does this population contain both monogamous and polyandrous groups simultaneously? What factors determine whether specific groups are monogamous or polyandrous? The data from this study population tentatively support the hypothesis that adults should mate monogramously only if they have nonreproductive helpers (usually older offspring) to help rear infants. Without helpers, the reproductive success of both males and females is hypothesized to be higher, on average, if they mate polyandrously than if they mate monogamously. The proposed benefits of polyandry to males and females differ quantitatively, but in both cases benefits stem from the help that males provide in rearing young. The following results support this hypothesis. (1) Lone pairs were never seen to attempt breeding, and calculations suggest that the costs of lactation and infant-carrying are too great for lone pairs to have a high probability of being able to raise twin offspring (the normal litter size). (2) Polyandrous males and nonreproductive offspring contributed substantially to infant care, particularly infant-carrying (fig. 2). (3) Adult males carried infants approximately twice as often as did lactating females, presumably because of the combined costs of (a) lactation (Fig. 3) and (b) infant-carrying (Fig. 4). The proximate causes of cooperative polyandry inS. fuscicollis appear to be different from those responsible in several bird species, showing that cooperative polyandry is a complex phenomenon.     

Perceived predation risk and mate defense jointly alter the outcome of territorial fights

Virtually all animal conflicts occur over access to mates or resources that affect survival, the two key components of fitness. In this paper, we report that predation risk and mate defense jointly affect the outcomes of contests between male sand crabs (Scopimera globosa) for burrows in which crabs mate and take shelter from predators. We observed the contests under three different conditions: (1) the natural condition of low predation risk and without the presence of a female; (2) the first experiment in which we imposed upon only intruding males the perception of predation risk—by digging them from their burrows, capturing and handling them, and placing them into other males’ burrows—to increase the value of the burrows for the intruders as shelter, and (3) the second experiment in which we repeated this treatment but increased the resource value of the burrow to the resident by placing a female in his burrow. The difference in body size between contestants was the main determinant of victory in all analyses. However, perceived predation risk also partly affected the outcomes of the fights: The motivated intruders were likely to win even when they were a little smaller than the residents. In addition, defense of a female had a significant effect on the outcomes of fights: The motivated residents won more fights than the motivated intruders, indicating that these two treatments caused asymmetric increases of the resource value. This is the first report of two external factors simultaneously raising resource value, affecting motivation of contestants, and altering the outcome of fights.     

Add-on conservation benefits of marine territorial user rights fishery policies in central Chile.

To combine the rational use of marine benthic resources and economic development of small-scale fishers, Chile passed legislation in 1991 establishing a comanagement policy that grants exclusive territorial user rights for fisheries (TURFs) to artisanal fisher organizations in well-defined inshore coastal areas, known as Management and Exploitation Areas for Benthic Resources (MEABRs). In general the policy has been proclaimed a management and economic success because benthic resource abundances have increased inside MEABRs in comparison with open-access areas. However, there is a lack of studies assessing the impact of this management policy on nontargeted subtidal species and community assemblages and the policy's implications for biodiversity and conservation. This study starts to fill this gap and links the allocation of TURFs for benthic resources with add-on conservation benefits for species that are not directly linked with the fishery policy. Comparative subtidal surveys inside vs. outside MEABRs were used to assess the effects of three MEABRs on managed targeted benthic species, biodiversity (species richness), and community assemblages in central Chile. Surveys focused exclusively on subtidal kelp forest habitats dominated by Lessonia trabeculata, spanning 4-12 m in depth and with similar levels of habitat complexity. The study comprised: (1) quantification of kelp forest complexity, (2) understory survey of sessile species, (3) quantification of conspicuous benthic macroinvertebrates, including those under management, and (4) quantification of reef-fish species inside the kelp habitat. Results showed population enhancement of target-managed invertebrates inside MEABRs. Moreover, reef-fish species were significantly more diverse and abundant inside MEABRs, and community assemblages of nontarget benthic invertebrates and reef fish were significantly different inside vs. outside MEABRs. The comanagement of inshore benthic resources in Chile, through MEABRs aims for the sustainability of invertebrate and algae stocks. However, our study shows that this management tool, which in practice restricts access to the entire management area, provides important conservation add-on effects for species that are not the focus of the management policies. Therefore, in Chile, the hundreds of already established MEABRs could represent an important ancillary network, which complements the biodiversity objectives of fully protected areas such as no-take marine protected areas or others.     

Costs and benefits of pocket gopher foraging: linking behavior and physiology

Animals can attain fitness benefits by maintaining a positive net energy balance, including costs of movement during resource acquisition and the profits from foraging. Subterranean rodent burrowing provides an excellent system in which to examine the effects of movement costs on foraging behavior because it is energetically expensive to excavate burrows. We used an individual-based modeling approach to study pocket gopher foraging and its relationship to digging cost, food abundance, and food distribution. We used a unique combination of an individual-based foraging-behavior model and an energetic model to assess survival, body mass dynamics, and burrow configurations. Our model revealed that even the extreme cost of digging is not as costly as it appears when compared to metabolic costs. Concentrating digging in the area where food was found, or area-restricted search (ARS), was the most energetically efficient digging strategy compared to a random strategy. Field data show that natural burrow configurations were more closely approximated by the animals we modeled using ARS compared to random diggers. By using behavior and simple physiological principles in our model, we were able to observe realistic body mass dynamics and recreate natural movement patterns.     

Costs and benefits of surplus offspring in the lesser kestrel (Falco naumanni )

Lesser kestrels (Falco naumanni) lay clutches which appear excessive as only 3% of them yield as many young as eggs laid. Four hypotheses may explain the adaptive value of producing surplus eggs: (1) the bet-hedging hypothesis assumes that the environment varies unpredictably and surplus eggs serve to track uncertain resources; (2) the ice-box hypothesis suggests that surplus offspring serve as a reserve food during a period of shortage; (3) the progeny choice hypothesis says that parents produce surplus offspring in order to choose these with higher fitness; and (4) the insurance-egg hypothesis proposes that extra eggs are an insurance against the failure of any egg. To test the significance of this strategy in the lesser kestrel, an experiment manipu-lating brood size at hatching was carried out over 2 years, with good and bad feeding conditions. The experiment consisted of adding a chick to experimental broods where one egg failed to hatch or removing a randomly selected chick from experimental broods where all eggs had hatched. Independently of annual food availability, pairs with brood sizes reduced by one chick fledged more nestlings than pairs with brood size equalling their clutch sizes. Body condition of young was also better in the former group, but only in 1993 (a high-food year). Independently of year, mean local survival of parents with complete broods at hatching was lower than for parents raising reduced broods. These results supported only the insurance-egg hypothesis which says that surplus eggs may be an insurance against the failure of any egg, but parents may suffer reproductive costs when all eggs hatch. Received: 17 January 1997 / Accepted after revision: 27 April 1997     

Genetic variation in defense chemistry in wild cabbages affects herbivores and their endoparasitoids

Populations of wild Brassica oleracea L. grow naturally along the Atlantic coastlines of the United Kingdom and France. Over a very small spatial scale (i.e., <15 km) these populations differ in the expression of the defensive compounds, glucosinolates (GS). Thus far, very few studies have examined interactions between genetically distinct populations of a wild plant species and associated consumers in a multitrophic framework. Here, we compared the development of a specialist (Pieris rapae) and a generalist (Mamestra brassicae) insect herbivore and their endoparasitoids (Cotesia rubecula and Microplitis mediator, respectively) on three wild populations and one cultivar of B. oleracea under controlled greenhouse conditions. Herbivore performance was differentially affected by the plant population on which they were reared. Plant population influenced only development time and pupal mass in P. rapae, whereas plant population also had a dramatic effect on survival of M. brassicae. Prolonged development time in P. rapae corresponded with high levels of the indole GS, neoglucobrassicin, whereas reduced survival in M. brassicae coincided with high levels of the aliphatic GS, gluconapin and sinigrin. The difference between the two species can be explained by the fact that the specialist P. rapae is adapted to feed on plants containing GS and has evolved an effective detoxification system against aliphatic GS. The different B. oleracea populations also affected development of the endoparasitoids. Differences in food-plant quality for the hosts were reflected in adult size in C. rubecula and survival in M. mediator, and further showed that parasitoid performance is also affected by herbivore diet.     

Intensity of territorial defense in red squirrels: an experimental test of the asymmetric war of attrition

Summary Red squirrels defend exclusive, individual territories year round, 20% to 50% of females do not breed in any given year, and breeding females raise juveniles on their territories. Breeding is asynchronous, and the offspring of early-breeding females are more likely to hold an independently won territory than are late-born offspring. Based on the asymmetric war of attrition, we made the following predictions: (1) squirrels would respond more intensely to the calls of unfamiliar individuals than to the calls of neighbors; (2) breeding females would respond more intensely to unknown calls than would non-breeding females or males; (3) early-breeding females would respond more intensely than would late-breeding females to unknown calls; and (4) all classes of squirrels would respond similarly to the calls of neighbors. Playback experiments supported the predictions. Alternative hypotheses of kin selection, risk of infanticide, and seasonal difference in intruder pressure could not explain the results.     

Costs and benefits of home range shifts among vervet monkeys (Cercopithecus aethiops) in Amboseli National Park,Kenya

Summary A decline in the density of trees and the deterioration of the habitat of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya, have been accompanied by movement of vervet groups into unfamiliar areas. These home range shifts were documented over a period of 26 months. The largest group moved into the home ranges of neighboring groups and acquired more trees, a resource that had become more limited. The groups whose home ranges were encroached upon did not share their home ranges with the intruders, but instead moved into areas that were unoccupied and had a lower density of trees. The largest group was thus more successful than smaller groups in competition for limited resources. For all but one group, movement into new areas was directly associated with the disappearances of female and immature monkeys. Most of the disappearances were attributable to predation. These results suggest that the cost of predation in unfamiliar areas may contribute to the evolution of philopatry in vervets. However, under unstable conditions, resource competition forces these animals to disperse (in groups) despite the high risk of predation. Offprint requests to: L.A. Isbell     

The evolution of marmot sociality: II. Costs and benefits of joint hibernation

Summary Social groups of alpine marmots (Marmota marmota) were studied for 7 years. The groups consisted of a territorial pair and up to 18 lower ranking animals of various ages, mostly the pair's offspring (Tables 1, 2). Group members lived in a common home range and always hibernated together in one hibernaculum. Groups with older, subordinate animals experienced slightly higher summer mortality but significantly reduced winter mortality (Fig. 1). Infant winter mortality was further decreased if most older subordinates were potentially their full sibs (Fig. 2). Subordinate group members lost less mass during winter with increasing size of the hibernating group, but this trend was reversed when infants were present. Furthermore, augmented mass loss due to low hibernaculum quality became evident (Fig. 3). Apart from these effects, the presence of infants caused additional mass loss in potential full sibs. The opposite was found in subordinates certainly descending from other parents than those of the infants (Table 3).Winter mortality and mass loss data revealed (i) a general benefit of joint hibernation, (ii) an unavoidable cost of infants' presence to other group members, (iii) that only potential full sibs helped in warming infants, (iv) that helping was energetically expensive and increased infant survival. The evolution of postponed dispersal in ground-dwelling squirrels has been attributed to the direct fitness gained by enhanced reproductive chances of offspring when not expelled from the natal territory (Armitage 1981, 1987, 1988). This study shows that group living in alpine marmots has benefits during winter and indicates the additional importance of kin selection in marmot social evolution.     

Costs and benefits of temporary brood desertion in a Neotropical harvestman (Arachnida: Opiliones)

Brood desertion is a life history strategy that allows parents to minimize costs related to parental care and increase their future fecundity. The harvestman Neosadocus maximus is an interesting model organism to study costs and benefits of temporary brood desertion because females abandon their clutches periodically and keep adding eggs to their clutches for some weeks. In this study, we tested if temporary brood desertion (a) imposes a cost to caring females by increasing the risk of egg predation and (b) offers a benefit to caring females by increasing fecundity as a result of increased foraging opportunities. With intensive field observations followed by a model selection approach, we showed that the proportion of consumed eggs was very low during the day and it was not influenced by the frequency of brood desertion. The proportion of consumed eggs was higher at night and it was negatively related to the frequency of brood desertion. However, frequent brood desertion did not result in higher fecundity, measured both as the number of eggs added to the current clutch and the probability of laying a second clutch over the course of the reproductive season. Considering that harvestmen are sensitive to dehydration, brood desertion during the day may attenuate the physiological stress of remaining exposed on the vegetation. Moreover, since brood desertion is higher during the day, when egg predation pressure is lower, caring females could be adjusting their maternal effort to the temporal variation in predation risk, which is regarded as the main cost of brood desertion in ectotherms.