Primate Conservation along the Tana River, Kenya: An Examination of the Forest Habitat

The Tana River National Primate Reserve, Kenya, was established in 1976 to preserve the endemic and endangered Tana River red colobus ( Colobus badius rufomitratus ) and crested mangabey ( Cercocebus galeritus galeritus ). Between 1975 and 1985 their populations declined by 80% and 45%, respectively. A study addressing primate-to-habitat relationships was conducted in 12 forest areas to determine what attributes appear most important to the in situ preservation of both endangered primates. Correlations among the abundances of primate groups, their group sizes, and the structural, resource, disturbance, and spatial attributes of the forest were used to distinguish the quality of forest patches. Intraforest habitat quality was examined, using canonical variate analysis to discriminate primate ranging patterns based on canopy and subcanopy tree composition. The colobus and mangabey show positive relationships to interior-forest habitat and appear susceptible to forest disturbances that reduce forest area or that increase forest edge and intraforest disturbance. Forest loss, fragmentation, and consequent reduction in the area-to-perimeter ratio of the remaining forests measured from 1960 to 1975 provide a partial explanation for the decline in primate populations. The results suggest that a combination of primary food items and seasonal food resources in large, high-stature, closed-canopy forests is the best predictor of high-quality habitat for these monkeys. Stewardship should be directed at the preservation of these areas or the restoration of forests toward this habitat model     

Maximization of foraging efficiency and resource defense by group retrieval in the ant Formica schaufussi

Summary Workers of the ant Formica schaufussi forage as individuals and cooperate in groups to retrieve arthropod prey. In 2 sample years, group-transported prey were on average 6.8 and 4.7 times heavier than individually retrieved items, and the average loading ratios of groups were greater than the loading ratios of single foragers. Retrieval group size was adjusted to prey size, and prey transport velocity for individuals and groups tended to decrease with increasing prey weight. The efficiency of individual and group retrieval, estimated from calculations of the prey delivery rate to the nest (PDR) achieved by each foraging mode, varied as a function of prey size. Individual retrieval maximized PDR at a prey weight of 19.5 mg, and group transport maximized PDR at 190 mg. Although the PDR maxima of an individual in a group and a solitary forager were approximately equal, depending on prey size, group transport may maximize foraging efficiency. Group transport also decreased interference competition from sympatric ant species. Group-transported prey having a greater likelihood of successful retrieval were within the size range of prey that maximized foraging efficiency. Transport group size appeared to be more important in prey defense than in increasing prey transport velocity, suggesting an important role of group size in competitive ability.Offprint requests to: J.F.A. Traniello     

All-offspring dispersal in a tropical mammal with resource defense polygyny

In polygynous mammals, males are usually responsible for gene flow while females are predominantly philopatric. However, there is evidence that in a few mammalian species female offspring may disperse to avoid breeding with their father when male tenure exceeds female age at maturity. We investigated offspring dispersal and local population structure in the Neotropical bat Lophostoma silvicolum. The mating system of this species is resource defense polygyny, with the resource being active termite nests, excavated by single males, which are then joined by females. We combined field observations of 14 harems during 3 years and data about the genetic structure within and between these groups, calculated with one mitochondrial locus and nine nuclear microsatellite loci. The results show that both male and female offspring disperse before maturity. In addition, we estimated life span of excavated termite nests and the duration they were occupied by the same male. Our findings suggest that long male tenure of up to 30 months is indeed a likely cause for the observed dispersal by female offspring that can reach maturity at a low age of 6 months. We suggest that dispersal by offspring of both sexes may occur quite frequently in polygynous tropical bats and thus generally may be more common in mammals than previously assumed.     

Sociality or territorial defense? The influence of resource renewal

Summary The rate with which resources in an area recover from local exploitation should influence the costs to an inhabitant of sharing it with neighbors. I develop a model which predicts the costs of tolerating conspecific foragers (or the benefits of excluding them) as a function of a predator's rate of harvesting prey and the prey's renewal rate. The predictions are consistent with patterns of social grouping observed in small African carnivores. A generalization of the model considers alternate forms of resource renewal (logistic, constant, or exponential) and suggests that not only the average rate of renewal but also the details of its time course should influence animal spacing patterns.     

Conspecific threat,predation avoidance,and resource defense: implications for grouping in langurs

Data on langur (Presbytis entellus) populations were gathered from the literature to test the importance of three selective pressures in determining group size and composition: predation pressure, intergroup resource defense, and conspecific threat. There were no detectable difference in the size of groups in populations facing nearly intact predator communities compared to those populations where predators were severely reduced in number or absent, although there was a trend for the number of adult males per group to increase in areas with nearly intact predator communities. Using population density as an indirect measure of the frequency of intrusions into a group’s home range and thereby as an index of the demographic pressure favoring resource defense, we predicted that higher densities would result in larger defensive coalitions and higher numbers of females per group. This prediction was not upheld. Our third selective pressure, conspecific threat, encompasses those selective forces resulting from physical attack on females, infants, and juveniles. Our index of conspecific threat uses the number of non-group males divided by the number of bisexual groups, because in langurs, the major source of conspecific threat derives from non-group males who, following group take-over, kill infants, wound females, and expel juveniles from groups. This index of conspecific threat was strongly related to the mean number of resident females, was weakly related to the mean group size, but was not related to the number of males in the group. In addition, as predicted, populations with a high index of conspecific threat had higher levels of juvenile expulsion. These analyses were corroborated by a simulation model which used a computer-generated series of null populations to calculate expected slopes of immatures regressed on adult females. These randomly generated populations, matched to means and ranges of real populations, allowed us to determine if deviations of the observed slopes from the expected null slopes could be explained by variation in predation pressure, population density, or conspecific threat. We found no evidence that predation pressure was associated with decreases in immature survival in smaller groups, as would be predicted by the predation-avoidance hypothesis. We found no evidence that immature survival was compromised by small group size in high-density populations, as would be predicted by the resource-defense hypothesis. However, as the index of conspecific threat increased, groups with larger numbers of females were more successful than groups with fewer females in reducing mortality or expulsion of immatures. Overall, conspecific threat received the strongest support as a selective pressure influencing langur group size and composition, suggesting that this selective pressure should be evaluated more widely as a factor influencing composition of animal groups. Received: 23 January 1995/Accepted after revision: 18 February 1996     

Individual variation in mate care by alpha males in crested tit winter flocks

Summary Winter flocks of crested tits Parus cristatus, typically consisting of two adults and one or two non-kin 1st-year birds, were observed to split up into subflocks in a way related to ambient temperature. On warm days, when they were foraging in subflocks without 1 st-year birds, alpha males always occupied the most preferred upper foraging sites, as expected from their top dominance rank. On cold days, when foraging in flocks with 1st-year birds, 8 out of 13 alpha males shifted to lower (less preferred) positions below their alpha mates while allowing the latter to forage at the best sites. As enhanced access to preferred microsites on days with high energy stress is believed to increase overwinter survival probability, this shifting behaviour of alpha males can be considered as a form of mate care. Out of 13 alpha males, however, 5 did not shift and always occupied the best foraging sites irrespective of flock composition. As (i) these non-shifting males were in poorer physical condition than shifting males, (ii) they scanned significantly less for predators than either females or shifting males when foraging in the uppermost tree parts, and (iii) four out of five non-shifting males were replaced by immigrants in early spring, absence of mate care during winter may be caused by constraints due to condition. High-quality-territory owners in poor condition at the end of autumn were most vulnerable to replacement by immigrants. Therefore, as four out of five replacements affected high-quality territories, selective intrusion by immigrants is suggested. Correspondence to: L. Lens     

Female mimicry and resource defense polygyny by males of a tropical rove beetle,Leistotrophus versicolor (Coleoptera : Staphylinidae)

Summary Adults of the staphylinid beetle Leistotrophus versicolor Grav. aggregate at vertebrate dung and carrion where males and females forage for adult Diptera. Some males aggressively exclude others from dung and carrion. Winners in male combat gain access to many females, which are often receptive at these foraging sites. The mating system can be categorized as resource defense polygyny. Males vary greatly in size, are larger than females on average, and have allometrically enlarged mandibles that they use in fighting. Large males consistently defeat smaller ones. Some males employ female mimicry in order to avoid aggression, remaining at dung where they forage and even obtain copulations while being courted by rival males. Female mimicry is most often practiced by males that are smaller than their rivals or by males that are unable to use their jaws aggressively because they are feeding or courting females when encountered by an opponent. Female mimicry is a conditional tactic of mature males; some individuals behave like females toward larger males but attack smaller rivals. Offprint requests to: J. Alcock     

Nest-building by crested tit Parus cristatus males: an analysis of costs and benefits

In a population of crested tits in Belgium, one out of three breeding males participated in nest-building. Since variation in the amount of nest-building among 11 building males was low, analysis of male nest-building allowed a logistic regression design. The probability of building increased with male condition and for late nests. The effect of male condition decreased with date, and date was the only factor predicting nest-building for repeat broods. The effect of male condition on nest-building suggests that building entails important costs for males. Based on correlative data, we argue that building by males shortens the interval between the start of nest-building and the onset of egg-laying by about 5 days. Since clutch size, incubation and nestling time did not differ between early and late broods, male nest-building apparently affects fledging date. Since early fledgers (early dispersers) are more likely to settle in future winter flocks, building behaviour by crested tit males should be favoured by natural selection. So far, male nest-building in hole-breeding birds might have been overlooked due to the widespread use of artificial nest-boxes in behavioural studies.     

Association of allozymes and temperature in the crested blenny Anoplarchus purpurescens

The crested blenny Anoplarchus purpurescens occurs intertidally in the Puget Sound area (USA), where it is subjected to substantial variation in temperature. A study of a polymorphic esterase in A. purpurescens has revealed a latitudinal cline and a correlation of allelic frequencies with summer water temperature. This finding agrees with previous results for a lactate dehydrogenase polymorphism in A. purpurescens, suggesting a pervasive genetic adaptation to temperature in this species. For each of these enzymes, comparisons within and between two species of Anoplarchus andicate an association between heterozygosity and temporal variability of the thermal environment.     

Aggressive brood defense by females in the stomatopod Gonodactylus bredini

Summary Defense behavior by brooding Gonodactylus bredini females was studied in staged fights against conspecific challengers. Females brooding eggs were more successful at defending their dwelling cavities than were nonreproductive females. Females caring for larvae were no more successful at cavity defense than were controls.Brooding females do not defend cavities by using more aggressive acts or by engaging in longer fights than nonreproductive females. The increase in defense success seems to be due to a greater willingness to continue a contest after being struck by a challenger. Also, brooding females make more frequent use of threats, in particular the Meral Spread display, than controls.Females defend cavities as vigorously as if they still had eggs 4 or 5 days after their eggs have been removed. Within the first 24 h after oviposition, a female's success at defending a cavity is greatly reduced compared to that of a female just 1 day later.     

Size matters: variation in the diet of chick and adult crested terns

We investigated ontogenetic, temporal and spatial patterns in the composition and size of prey in the diet of crested terns, Sterna bergii. Diet analyses indicated that crested terns are a generalist predator on surface-schooling clupeids (Australian anchovy Engraulis australis, sardine Sardinops sagax and blue sprat Spratelloides robustus), Degens leatherjacket Thamnaconus degeni, southern sea garfish Hyporhamphus melanochir, Australian herring Arripis georgianus, slender bullseye Parapriacanthus elongatus and barracouta Thyrsites atun. Ontogenetic differences in prey size indicated that adults are constrained in their foraging behaviour during the early chick-provisioning period by the need to self feed and select smaller prey that can be ingested by their chicks. Chicks consumed significantly higher proportions of clupeids than adults, which consumed mainly Degens leatherjackets and barracouta, suggesting that adults may select higher quality prey for their chicks compared to what they consume themselves. Spatial differences in prey composition were driven by differing proportions of sardine, Australian anchovy and Degens leatherjacket and could reflect local differences in the abundances of these prey. The size of prey taxa consumed by adults also reflected a North–South gradient in prey size. The large component of juvenile sardine in the diet of crested terns suggests future dietary measures may inform fisheries managers about changes in local juvenile sardine abundance. These data could assist in highlighting any fishery-related decreases in sardine recruitment and help ensure commercial fishing practices address principals of Ecologically Sustainable Development developed for Australian fisheries.     

On the extraction of an exhaustible resource by a monopoly

Hotelling's r-percent rule does not hold for monopoly extractors of durable exhaustible resources. An example with a nondurable resource in which the rule also fails to hold is presented. An economy with a fixed average propensity to save is modelled. The monopoly extractor recognizes that resource extraction, by affecting output and hence capital accumulation, affects future demand. The firm exploits this effect by causing the marginal profitability of extraction to grow faster or slower than the rate of interest, depending upon initial conditions. Conditions are developed under which the growth rate will be less than the interest rate.     

Resource defense polygyny shifts to female defense polygyny over the course of the reproductive season of a Neotropical harvestman

Although studies classify the polygynous mating system of a given species into female defense polygyny (FDP) or resource defense polygyny (RDP), the boundary between these two categories is often slight. Males of some species may even shift between these two types of polygyny in response to temporal variation in social and environmental conditions. Here, we examine the mating system of the Neotropical harvestman Acutisoma proximum and, in order to assess if mate acquisition in males corresponds to FDP or RDP, we tested four contrasting predictions derived from the mating system theory. At the beginning of the reproductive season, males fight with other males for the possession of territories on the vegetation where females will later oviposit, as expected in RDP. Females present a marked preference for specific host plant species, and males establish their territories in areas where these host plants are specially abundant, which is also expected in RDP. Later in the reproductive season, males reduce their patrolling activity and focus on defending individual females that are ovipositing inside their territories, as what occurs in FDP. This is the first described case of an arachnid that exhibits a shift in mating system over the reproductive season, revealing that we should be cautious when defining the mating system of a species based on few observations concentrated in a brief period. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Optimal defense: snails avoid reproductive parts of the lichen Lobaria scrobiculata due to internal defense allocation

The optimal defense theory (ODT) deals with defensive compounds improving fitness of a particular organism. It predicts that these compounds are allocated in proportion to the risk for a specific plant tissue being attacked and this tissue's value for plant fitness. As the benefit of defense cannot easily be measured in plants, the empirical evidence for ODT is limited. However, lichens are unique in the sense that their carbon-based secondary compounds can nondestructively be removed or reduced in concentration by acetone rinsing. By using such an extraction protocol, which is lethal to plants, we have tested the ODT by studying lichens instead of plants as photosynthetically active organisms. Prior to acetone rinsing, we found five times higher concentration of meta-scrobiculin in the reproductive parts (soralia) of Lobaria scrobiculata compared to somatic parts of this foliose epiphytic lichen species. At this stage, the lichen-feeding snail Cochlodina laminata avoided the soralia. However, after removal of secondary compounds, the snail instead preferred the soralia. In this way, we have successfully shown that grazing pattern inversely reflects the partitioning of the secondary compounds that have a documented deterring effect. Thus our study provides strong and novel evidence for the ODT.     

Unequal resource allocation among colonies produced by fission in the ant Cataglyphis cursor

How organisms allocate limited resources to reproduction is critical to their fitness. The size and number of offspring produced have been the focus of many studies. Offspring size affects survival and growth and determines offspring number in the many species where there is a trade-off between size and number. Many social insects reproduce by colony fission, whereby young queens and accompanying workers split off from a colony to form new colonies. The size of a new colony (number of workers) is set at the time of the split, and this may allow fine tuning size to local conditions. Despite the prevalence of colony fission and the ecological importance of social insects, little is known of colony fission except in honey bees. We studied colony fission in the ant Cataglyphis cursor. For clarity, "colony" and "nest" refer to colonies before and after colony fission, respectively (i.e., each colony fissions into several nests). The reproductive effort of colonies was highly variable: Colonies that fissioned varied markedly in size, and many colonies that did not fission were as large as some of the fissioning colonies. The mother queen was replaced in half of the fissioning colonies, which produced 4.0 +/- 1.3 (mean +/- SD) nests of markedly varied size. Larger fissioning colonies produced larger nests but did not produce more nests, and resource allocation among nests was highly biased. When a colony produced several nests and the mother queen was not replaced, the nest containing the mother queen was larger than nests with a young queen. These results show that the pattern of resource allocation differs between C. cursor and honey bees. They also suggest that C. cursor may follow a bet-hedging strategy with regard to both the colony size at which fission occurs and the partitioning of resources among nests. In addition, colony fission may be influenced by the age and/or condition of the mother queen, and the fact that workers allocating resources among nests have incomplete knowledge of the size and number of nests produced. These results show that the process of colony fission is more diverse than currently acknowledged and that studies of additional species are needed.     

Influence of dung volatiles on the process of resource selection by coprophagous beetles

Summary. Most dung beetles colonize the faeces of several vertebrate species without much discrimination, and are thus often considered as polyphagous. Recent studies have provided evidence for clear feeding preferences in scarab beetles colonizing dung of herbivore species, but little is known about these insects’ abilities to discriminate among odours from faeces of various herbivores. In this study, trophic preferences were examined using blocks of pitfall traps baited with dung from four different herbivore species, i.e., sheep, cattle, horse, and red deer, in a mountainous area of south-central France. 4941 coprophagous scarabs, belonging to 27 species, were captured. Beetles were more attracted to dung of sheep (2257 individuals) than that of cattle (1294 individuals), followed by deer dung (768 individuals) and horse dung (622 individuals). Eleven of the 27 beetle species collected had significant feeding preferences for one of the four dung types. For each insect species, trophic habits did not vary between the two different sites of trapping, an open pasture and a wooded habitat. In laboratory olfactometer bioassays, scarab beetles orientated preferentially towards the dung volatiles from the dung type they preferred in the field. Trypocopris pyrenaeus, Anoplotrupes stercorosus, and Aphodius rufipes were more attracted to volatile compounds from sheep dung, Onthophagus fracticornis significantly preferred horse dung volatiles, and Aphodius haemorrhoidalis responded positively to deer dung odours. The role of dung olfactory cues in the process of resource selection by dung beetles is discussed.     

Chemical defense of the ladybird beetle Epilachna paenulata

Summary. The defensive chemistry of the ladybird beetle Epilachna paenulata (Coleoptera: Coccinellidae) was characterized as a mixture of piperidine, homotropane and pyrrolidine alkaloids. Whole body extracts of adult beetles contain four major alkaloids: 1-(6-Methyl-2,3,4,5-tetrahydro-pyridin-2-yl)-propan-2-one; 1-(6-methyl-2-piperidyl)-propan-2-one; 9-aza-1-methyl-bicyclo[3.3.1]nonan-3-one and 1- (2′′- hydroxyethyl)-2-(12′-aminotridecyl)-pyrrolidine. Comparative studies of the defensive chemistry of eggs, larvae, pupae and adults showed both qualitative and quantitative differences in alkaloid composition among the four life stages, and also within adult age. Laboratory predation bioassays with wolf spiders showed that the adults are better protected than the larvae and pupae. Field tests showed the adult alkaloid extract to be deterrent to ants.