Life History of Cercopithecus mitis stuhlmanni in the Kakamega Forest, Kenya

Comparative data from wild populations are necessary to understand the evolution of primate life history strategies. We present demographic data from a 29-yr longitudinal study of 8 groups of individually recognized wild blue monkeys (Cercopithecus mitis stuhlmanni). We provide estimates of life history variables and a life table for females. Most females had their first infant at 7 yr. The mean interbirth interval was 28 mo, and decreased from 31 to 18 mo if the first infant died within a year. Interbirth intervals did not differ according to infant sex, but females had longer intervals after their first vs. subsequent births. Infant mortality was 23% and did not differ strongly by sex or mother’s parity. Maximal female lifespan was 32.5–34.5 yr. Across the lifespan, both survivorship and fecundity showed typical primate patterns. Survivorship was lowest in infants, leveled off among juveniles, and then decreased gradually with increasing age in later life. Fecundity was highest among young females and decreased among older females. Births were seasonal, with 64% occurring within 3 mo at the end of the dry season and beginning of the wet season. Survival to 12 mo was higher for infants born during drier months. Birth season timing is plausibly related to thermoregulation of infants, weanling foods, or maternal energy demand. Blue monkeys are a forest-dependent species with a very slow life history and relatively low immature and adult mortality rates compared to closely related guenons living in open habitats. Even among cercopithecines as a whole, they appear to have an exceptionally slow life history relative to body size. Differences in life history “speed” between blue monkeys and their close relatives seem to be related to lower juvenile and adult mortality in forests relative to more open habitats.     

Assessment of Threat Status and Management Effectiveness in Kakamega Forest, Kenya

To counteract an increasing biodiversity decline, parks and protected areas have been established worldwide. However, many parks lack adequate management to address environmental degradation. To improve management strategies simple tools are needed for an assessment of human impact and management effectiveness of protected areas. This study quantifies the current threats in the heavily fragmented and degraded tropical rainforest of Kakamega, western Kenya. We recorded seven disturbance parameters at 22 sites in differently managed and protected areas of Kakamega Forest. Our data indicate a high level of human impact throughout the forest with illegal logging being most widespread. Furthermore, logging levels appear to reflect management history and effectiveness. From 1933 to 1986, Kakamega Forest was under management by the Forest Department and the number of trees logged more than 20 years ago was equally high at all sites. Since 1986, management of Kakamega Forest has been under two different organizations, i.e. Forest Department and Kenya Wildlife Service. The number of trees logged illegally in the last 20 years was significantly lower at sites managed by the Kenya Wildlife Service. Finally, logging was lower within highly protected National and Nature Reserves as compared to high logging within the less protected Forest Reserves. Reflecting management effectiveness as well as protection status in Kakamega Forest, logging might therefore provide a valuable quantitative indicator for human disturbance and thus an important tool for conservation managers. Logging might be a valuable indicator for other protected areas, too, however, other human impact such as e.g. hunting might also prove to be a potential indicator.     

Habitat disturbance, severity and patterns of abundance in Kakamega Forest, Western Kenya

Patterns of plant species abundance were studied for five old patches in the Kakamega Forest in Western Kenya. All the five patches had been subjected to disturbances involving removal of top soil. Species diversity, species abundance, damage to individuals in various patches, quality of growth and distribution of life-forms differed among patches. The pattern of species abundance showed the most striking variation in conforming to various species dominance curves. Explanations for these differences in species quality appear to reflect the quality of the patches.

Résumé

On a étudié dans 5 anciennes parcelles de la Kakamega Forest à l'ouest du Kenya les schémas d'abondance en espèes végétales. Les 5 parcelles avaient été soumises a des perturbations comprenant l'évacuation de la couche supérieure du sol. La diversité des espèces, leur abondance, les dommages causés aux plantes indivi-duelles dans les différentes parcelles, le type de croissance et la distribution des espèces différaient selon les parcelles. Le schéma d'abondance des espèces montrait la variation la plus frappante par sa conformité aux courbes de dominance des différentes espèces. Les explications de ces différences de la qualité des espèces semble refléter la qualité des parcelles.     

Long-term tree population dynamics and their implications for the conservation of the Kakamega Forest, Kenya

Long-term studies of tree population dynamics play an important role in identifying the conservation needs of tropical forest ecosystems. We examined changes in tree population structure and composition over an 18-year period (1981–1999) in three plots located at the center of the Isecheno study site in the Kakamega Forest, Kenya, a forest with a history of logging and other anthropogenic disturbance. DBH size class distribution took the shape of an inverse J curve in both 1981 and 1999 and did not differ significantly between the two study periods. Stem density increased significantly during the study with most of the increase occurring in the smallest stem size class (10–14 cm DBH). Nearly all of the most common species in 1981 remained among the most common in 1999, though the density of pioneer species decreased by 21% during the study. Our results suggest that forest in the study plots remained relatively undisturbed and in good condition over the study period. Forest in the plots also appeared still to be recovering from the selective logging of large trees that took place at Isecheno in the 1940s. In addition to our longitudinal study, we compared tree population parameters at three additional Isecheno sites spread over a distance of 1 km that have experienced different histories of disturbance: (i) a lightly human disturbed site (LHD), (ii) a heavily human disturbed site (HHD), and (iii) a cattle disturbed site (CD). While all three sites were selectively logged in the 1940s, the main signs of disturbance today are footpaths at the LHD site, tree stumps at the HHD site, and wide cattle paths at the CD site. Not surprisingly, of the disturbed sites, the LHD site was in the best condition. Trees at the HHD site exhibited extremely poor recruitment into the small size classes, a condition that can probably be attributed to human exploitation of small trees for poles. The CD site appeared to be at an earlier successional stage than the other disturbed sites with its low mean DBH, high overall stem density, and high pioneer species stem density. Browsing and trampling of vegetation by cattle may be the source of the light gaps that have led to the abundance of pioneer species at this site. We conclude that conservation measures applied to central Isecheno, including the establishment of a forest station nearby and ranger patrols, appear to have succeeded, but that the prognosis for the Kakamega Forest in general is bleak if protection efforts are not increased in other parts of the forest, where anthropogenic disturbance remains high. We also note the considerable variation in tree population structure and composition that can occur within a small area depending on the local history of disturbance.     

Feeding Ecology of Guerezas in the Kakamega Forest,Kenya: The Importance of Moraceae Fruit in Their Diet

Eastern black-and-white colobus (Colobus guereza), or guerezas, have long been considered to be one of the most folivorous primates. I conducted a study of the feeding ecology of two guereza groups (T and O) over an annual cycle in the Kakamega Forest of western Kenya. I found that the annual diets of both groups comprised mostly of leaves (T: 48%, O: 57%) though fruit (T: 44%, O: 33%) also accounted for a substantial portion of the diet. In the six months when fruit was most abundant, fruit consumption constituted an average of 58% of T-group's monthly diet and 42% of O-group's monthly diet. In contrast to most previous studies of colobines, in which seeds were the primary fruit item consumed, almost all of the fruit eaten by guerezas at Kakamega consisted of whole fruits. At least 72% of the whole fruits consumed by T-and O-groups were whole fruits from trees in the Moraceae family, which dominates the tree family biomass at Kakamega. Unlike at sites where guerezas consumed fruit primarily when young leaves were scarce, at Kakamega guerezas ate fruit in accordance with its availability and irrespective of the availability of young leaves. My findings demonstrate that guerezas are more dietarily flexible than was previously known, which may help to explain why the species can survive in such a wide variety of forested habitats across equatorial Africa.     

Use of sleeping trees by black and white Colobus monkeys (Colobus guereza) in the Kakamega Forest,Kenya

Groups of black and white colobus monkeys, or guerezas (Colobus guereza), observed in the Kakamega Forest, Kenya, had weak fidelity for sleeping sites. Groups often slept in trees near commonly used food sources, which might reduce the time and energetic costs of travel. Although the home range of each group overlapped with four to seven others, groups seemed to avoid sleeping near other groups, which would give them immediate and exclusive access to nearby food sources in the morning. The number of times a species of tree was slept in was positively correlated with its density. This may have occurred because so many suitable sites were available that proximity to feeding trees could be obtained whether or not groups slept in the feeding trees. Groups slept in tall trees, which provide stable sleeping sites and which may provide protection from both aerial and ground predators. Groups were more tightly clustered on nights with greater visibility, which might reduce the risk of predation. Am. J. Primatol. 45:281–290, 1998. © 1998 Wiley-Liss, Inc.     

Mating Behavior, Sexual Selection, and Copulatory Courtship in a Promiscuous Beetle

The function of male movements during copulation is unclear. These movements may be a result of the necessary mechanics of insemination, or they may also have further function, for instance, stimulating or courting a female during mating, perhaps influencing female mate choice. We present data from three experiments exploring the mating behavior and copulatory movements of the highly promiscuous beetle Psilothrix viridicoeruleus. Male mating success in the struggle over mating was not related to male or female size (measured by weight) but successful males were more vigorous in terms of copulatory movements. These males took longer to mount females but copulated longer and remained mounted longer. We discuss these results in terms of the mating system of Psilothrix and also in terms of observations of the timing of insemination during copulation. We suggest that copulatory movements in this species are best understood as copulatory courtship.     

Conservation of biodiversity in the Arabuko Sokoke Forest, Kenya

Using an economic approach to provide a rationale for rainforestconservation has been a popular exercise in recent years. This paper uses suchan approach to assess the net value of the Arabuko Sokoke Forest in Kenya. Theeconomic benefits associated with the forest derived by local and globalpopulations are estimated by combining evidence from existing studies and theresults of a contingent valuation study carried out by the authors. Thesebenefits are set against the cost of preserving the forest to the Kenyan ForestDepartment. Even when the opportunity cost of the forest land is omitted fromthe costs of forest preservation, and when the revenues generated from theGlobal Environment Facility (GEF) funded project are included, the costs offorest conservation outweigh the benefits. It is only when non-use andexistencevalues are included (which are not realised by the Kenyan population) that theforest benefits exceed the costs. The paper concludes by arguing that, althoughsome projects within the Arabuko Sokoke Forest have been successful incapturingsome of the economic value associated with the forest, more needs to be done todesign additional capture mechanisms so that a greater proportion of the globalbenefit of the forest can be realised by local populations and localgovernments.     

Infanticide in <Emphasis Type="Italic">Cercopithecus mitis stuhlmanni</Emphasis> in the Kakamega Forest,Kenya: Variation in the Occurrence of an Adaptive Behavior

Infanticide by males is widespread across mammals and especially prevalent among primates. Considerable research has examined how fitness benefits can explain the occurrence of this behavior; less is known, however, about intrapopulation variation in its occurrence. We evaluated 10 infanticides by males in wild blue monkeys according to the sexual selection hypothesis. To explore intrapopulation variation in occurrence of infanticide, we compared these cases to 38 cases that were contextually similar but in which infanticide did not occur. We examined male reproductive benefit, infant age, maternal parity, postconception estrus, group defense, available mating partners, and context of takeover. We based comparisons on daily or near daily records of male presence in the study groups, infant birth dates, and male-female sexual interactions. Infanticides followed predictions of the sexual selection hypothesis: males were unlikely to kill their own offspring, the period for the mother’s return to conception was reduced by half, and males increased their chance of siring her next offspring. Difference in male reproductive benefit, costs, and motivation did not fully explain the observed variation in infanticide occurrence. Infants were more likely to be spared if they were older when a male first arrived, or if their mother had mated with the male in the second month after conception. The most important determinant of infant fate, however, was male identity, a finding consistent with 2 scenarios: 1) an infanticidal tendency may be influenced by a genetic polymorphism that is not fixed in this population or 2) infanticidal behavior may be a conditional male strategy. Further research on intrapopulation variation in infanticidal behavior should focus especially on characteristics of males.     

Male Demography,Female Mating Behavior,and Infanticide in Wild Patas Monkeys (Erythrocebus patas)

Infanticide by males has been hypothesized to be a naturally selected behavioral strategy that increases the infanticidal male's reproductive success. The sexual selection hypothesis has been challenged via alternative, nonadaptive hypotheses that dispute its empirical and theoretical bases. Two of the most widely recognized alternatives are the social pathology hypothesis, in which infanticide results from overcrowding or recent human disturbance, and the generalized aggression hypothesis, in which infanticide is an epiphenomenon of increased male aggression. We report the first case of infanticide in wild, seasonally breeding patas monkeys (Erythrocebus patas) living at a low population density in a stable habitat, conditions which do not support the social pathology hypothesis. Its exceptional occurrence is consistent with the sexual selection hypothesis: over a 7-year period the infanticidal male was the only one of 13 resident males that was not present during the actual conception season but was present during the following birth season. Also consistent with this hypothesis, mothers were differentially targeted for male aggression, which increased sevenfold during the days surrounding the infanticide and then decreased to baseline levels after the infanticide. Aggression targeted at mothers does not support the generalized aggression hypothesis. As predicted by the sexual selection hypothesis, females began soliciting mating immediately after the infanticide, despite its occurrence in the nonconceptive season.     

Influence of Plant and Soil Chemistry on Food Selection,Ranging Patterns,and Biomass of <Emphasis Type="Italic">Colobus guereza</Emphasis> in Kakamega Forest,Kenya

Nutritional factors are among the most important influences on primate food choice. We examined the influence of macronutrients, minerals, and secondary compounds on leaf choices by members of a foli-frugivorous population of eastern black-and-white colobus—or guerezas (Colobus guereza)—inhabiting the Kakamega Forest, Kenya. Macronutrients exerted a complex influence on guereza leaf choice at Kakamega. At a broad level, protein content was the primary factor determining whether or not guerezas consumed specific leaf items, with eaten leaves at or above a protein threshold of ca. 14% dry matter. However, a finer grade analysis considering the selection ratios of only items eaten revealed that fiber played a much greater role than protein in influencing the rates at which different items were eaten relative to their abundance in the forest. Most minerals did not appear to influence leaf choice, though guerezas did exhibit strong selectivity for leaves rich in zinc. Guerezas avoided most leaves high in secondary compounds, though their top food item (Prunus africana mature leaves) contained some of the highest condensed tannin concentrations of any leaves in their diet. Kakamega guerezas periodically traveled great distances to exploit rare foods (bark from exotic Myrtaceae trees and soil) outside their normal home ranges. Our results suggest that these journeys were driven by the fact that these rare foods contained exceptionally high sodium concentrations, a mineral believed to be deficient in the guereza's usual diet. Lastly, our results are consistent with the pattern established across other Paleotropical rain forests in which colobine biomass can be predicted by the protein-to-fiber ratio in mature leaves. Of the 8 rain forests for which the relevant data are available, Kakamega features the second highest mature leaf protein-to-fiber ratio as well as the second highest colobine biomass.     

Activity and Ranging Patterns of Guerezas in the Kakamega Forest: Intergroup Variation and Implications for Intragroup Feeding Competition

From March 1997 to February 1998, I investigated the activity patterns of 2 groups and the ranging patterns of 5 groups of eastern black-and-white colobus (Colobus guereza), aka guerezas, in the Kakamega Forest, Kenya. Guerezas at Kakamega spent more of their time resting than any other population of colobine monkeys studied to date. In addition, I recorded not one instance of intragroup aggression in 16,710 activity scan samples, providing preliminary evidence that intragroup contest competition may be rare or absent among guerezas at Kakamega. Mean daily path lengths ranged from 450 to 734 m, and home range area ranged from 12 to 20 ha, though home range area may have been underestimated for several of the study groups. Home range overlap was extensive with 49–83% of each group's range overlapped by the ranges of other groups. Despite the high level of home range overlap, the frequently entered areas (quadrats entered on 30% of a group's total study days) of any one group were not frequently entered by any other study group. Mean daily path length is not significantly correlated with levels of availability or consumption of any plant part item. Mean daily path length is also not significantly correlated with group size, though the largest group did have the longest mean daily path length. This finding suggests that intragroup scramble competition may have been rare or absent among guerezas at Kakamega except perhaps in the largest group, which was unusually large.     

Wealth, Status, and Reproductive Success among the Mukogodo of Kenya

The evolutionary biological hypothesis that culturally defined values and goals are proximate means of enhancing reproductive success is tested on data from the Mukogodo, a small group of Maa-speaking pastoralists in north-central Kenya who value the accumulation of livestock. The results support the prediction that, at least among males, livestock wealth should correlate with reproductive success. This correlation appears to be due mainly to greater polygyny among wealthier men. Lower age at first marriage among wealthier men may also contribute to the correlation between livestock wealth and reproductive success. The association between livestock wealth and reproductive success does not appear to be due to the productivity of wives and children, to bridewealths obtained when daughters marry, or to the effects of wealth on the reproductive success of men's wives.     

Livestock production,age, and gender among the Keiyo of Kenya

This article is based on a study of time allocation and decision making patterns among the Keiyo in three ecological zones of Elgeyo-Marakvet District, Kenya. It finds that age and gender are important factors to consider when examining livestock production among African smallholder farmers. This is especially true where males are absent from home for wage labor. In these situations women, young males between the ages of 6–15, and persons of both sexes aged 65 and over spend considerable amounts of time engaged in livestock-related activities. These women are also more actively involved in decision making concerning the marketing of milk. The results indicate that there is a definite need to consider the contributions of these other persons when efforts are being made to improve elements of the livestock sector.     

Distribution and population size of the threatened East Coast Akalat in Arabuko-Sokoke Forest,Kenya

Matiku, P.M., Bennun, L.A. & Nemeth, E. 2000. Distribution and population size of the threatened East Coast Akalat in Arabuko-Sokoke Forest, Kenya. Ostrich 71 (1 & 2): 282–285.

The East Coast Akalat Sheppardia gunningi sokokensis is one of six globally threatened bird species that occur in the Arabuko-Sokoke Forest, the largest remnant of forest on the East African coast. The akalat has been little studied but appears to have a patchy distribution both between and within forests. From October 1995 to February 1996 we studied the akalat's status in Arabuko-Sokoke, carrying out transect counts and plot-based surveys to determine its distribution and population size. Playback of calls was used to stimulate the bird to reveal its presence. The akalat was present in two of the forest's main vegetation types, Cynometm woodland and Mixed forest. It was absent from the more open Brachystegia woodland, and from low-canopied intermediate Cynometra and Cynometra thicket. In Cynometra woodland, the akalats were evenly distributed at high densities (estimated at 87 pairs/km²). Within the Mixed forest distribution was patchy, and the akalats were absent from areas with high human disturbance. Where the species did occur, densities were lower (estimated at 12 pairs/km²) than in the Cynometra woodland. Arabuko-Sokoke may hold up to 9 000 pairs of this species, perhaps the largest single population in the world. Most of these are in the Cynometra woodland, which is also of crucial conservation importance for the threatened Sokoke Scops Owl Otus ireneae.     

Ranging Behavior of Proboscis Monkeys in a Riverine Forest with Special Reference to Ranging in Inland Forest

We observed a unimale group (BE-Group) of proboscis monkeys (Nasalis larvatus) comprising an α-male, 6 adult females, and several immatures from May 2005 to May 2006. We followed the group for 2014 h along the Menanggul River, Sabah, Malaysia (118°30′E, 5°30′N). Observations focused mainly on ranging behavior. We determined availability and seasonal changes in plant species consumed by the members of the group by vegetation surveys in a 2.15-ha area along 200–500 m trails in the riverine forest. During the observation period, the group ranged ≤800 m from the riverbank, within a total range of 138.3 ha. The daily path length of the group ranged from 220 to 1734 m (mean, 799 m), and daily path length correlates negatively with fruit availability. The monkeys were apt to remain within a small range in fruit-abundant seasons. Because the monkeys preferred to feed on fruits of dominant plant species in the study area, their daily path length may decrease on days when they feed on fruits. The core areas of the group’s home range were along the river because the monkeys typically returned to riverside trees to sleep. The group most often used areas that were nearer the riverbank and where the availability of fruits was higher. The most frequently used grids were the ones where the group often had sleeping sites and crossed the river. Avoiding predation may be the main reason for river crossing and selecting particular sleeping sites; hence not only food availability but also the risk of predation appears to influence the ranging of the BE-Group.