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The Impact of Tourism on Ring-Tailed Lemur (Lemur catta) Behaviour, Home Range Size and Habitat Use at Berenty Reserve, Madagascar
Abstract
The ring-tailed lemur (Lemur catta), endemic to Madagascar, is considered 'Near Threatened' by the International Union of Conservation of Nature. The limited numbers are the result of anthropogenic activities. No previous research has specifically investigated the possible impact of primate tourism on free-ranging ring-tailed lemurs; which is imperative to understanding future conservation measures that need to be enforced. This study was carried out between February and April at Berenty Reserve in Madagascar, and investigated the impact of tourist pressure on the behaviour, home range size and habitat usage of two troops of ring-tailed lemur subjected to different intensities of tourism. Tourist presence was found to negatively influence the activity budget behaviour of resting, and positively influence intra-troop aggression. Furthermore, increasing tourist density was found to negatively influence feeding on human food resources. Greater rates of scent-marking (both a hypothesised self-directed behaviour indicative of anxiety, as well as an activity of 'tradition' in ring-tailed lemurs) were apparent when tourists were present. Increased inter-troop aggression was found to be positively related to greater rates of scent-marking, tourist feeding interactions, and feeding on human food resources. Minimum Convex Polygons and Kernel Density Estimators were used to analyse the home ranges, revealing that food resource distribution influenced home range size and usage. This research suggested that highly clumped and nutritious provisioned foods (including introduced tree species), resulted in greater rates of intra-troop and inter-troop aggression (most notably amongst females) and scent-marking behaviours. It was concluded that overcrowding, and smaller home ranges of troops found at the tourist front are indirectly due to tourist pressure. This study therefore highlights the need for improved monitoring of food provisioning and ways to control and mitigate the overcrowding lemur populations.
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At Berenty Reserve, Madagascar, we have set up a main study area of 14.2 ha to study the demographic changes of ring-tailed lemurs for 11 years from 1989 to 2000. We also have set up a broader study area of 30.4 ha to study the environmental changes, including the main study area. In the main study area, due to the social changes such as troop divisions and evictions, the number of troops increased from three to six. Consequently, home range size of a troop decreased. In the broader study area, there were 475 large trees belonging to 14 species and 9 families. The most abundant species was kily (Tamarindus indica) (n=289), the second benono (Acacia rovumae) (n=74), the third voleli (Nestina isoneura) (n=66). These three species accounted for 90.3 % of all large trees. Population density of large trees was 15.6 per ha, and that of idly trees was 10.3 per ha. In 1989, 12.7 kily trees existed per ha within the main study area of 14.2 ha.In 2000, we re-measured the size of kily trees. Within the main study area of 14.2 ha, the density of idly trees decreased to 11.2 per ha. This decrease was due to the number of dead kily trees exceeded the number of newcomers. In contrast, the number of ring-tailed lemurs over one-year old increased from 63 in 1989 to 89 in 2000. As a result, the number of kily trees per animal was decreased from 2.8 to 1.8. Among troops, there was a great variation in the number of idly trees per animal. Troop CX occupied the richest area (4.7 kily trees/animal), and Troop T2 ranged the poorest area (0.6 kily trees/animal). On the other hand, the amount of kily fruits may fluctuate by each individual tree, area, and year. For example, the harvest of idly fruits was very poor in 2000, and the abundance score of kily fruits for Troop CX was lower than average score of the main study area. The number of brown lemurs (Eulemur fulvus) was also increasing. Since brown lemurs' feeding habits were very similar to ring-tailed lemurs, it is likely that both the within species and inter-species competitions are becoming intense for ring-tailed lemurs.
Primary forest cover in western Madagascar declined from 12.5% in 1950 to 2.8% in 1990. Approximately 23% of remaining forest is located within reserves, but this is no guarantee of protection. Forest cover in the Andranomena Reserve in western Madagascar has been reduced by 44% since 1950. The identification of priority areas for establishment of development projects has been constrained by lack of information on the distribution, abundance, and habitat requirements of threatened species and the size, condition, and threats to survival of forest remnants. We evaluate a rapid fauna survey and habitat modeling procedure specifically designed to generate information for reserve selection and design using Geographical Information Systems (GIS). The study was conducted in the largest remnant (94,000 ha) of primary monsoon rainforest in western Madagascar, using lemurs as an indicator group. Lemur abundance and microhabitat variables were measured at 64 stratified survey sites dispersed throughout the region. These records were incorporated into a GIS with other environmental and landuse data derived from maps and satellite imagery. Statistical procedures were applied to analyze species habitats, predict and map species distributions, and estimate species population sizes throughout the study area. Patterns of slash and burn agriculture and anthropogenic disturbance were also analyzed to identify areas at low risk of cultural disturbance. A map of disturbance risk was overlaid with a map of relative lemur biodiversity to identify conservation priority areas with high biodiversity and low risk of short term disturbance. Lemur diversity was most strongly influenced by habitat clearing and human disturbance (harvesting, stock grazing, and hunting) within 8 km of villages. Lemur diversity was highest in a region of higher elevation and rainfall, distant from villages and roads. The only existing reserve in the study area was located in a region of high disturbance risk. Almost half the reserve has been degraded during the past 45 years. There is considerable scope for re-allocation of land use within the study area to include a core protected area, a buffer zone with controlled hunting and timber harvesting, and an adjacent ecotourism facility. We found that information generated by a stratified biodiversity and landuse survey at a sampling intensity of less than 0.1% was sufficient to provide an objective foundation for regional biodiversity planning.
La cobertura de bosques primarios en el oeste de Madagascar disminuyó de 12.5% en 1950 a 2.8% en 1990. Aproximadamente un 23% del bosque remanente está localizado en reservas, pero esto no garantiza su protección. Las reservas están sujetas a caza, recolección, pastoreo y actividades de agricultura con tumba y quema. La cobertura de la reserva de Andranomena en Madagascar ha sido reducida en un 44% desde 1950. En vista de esta amenaza, los bosques tropicales estacionales deciduos de monson del oeste de Madagascar pueden ser considerados entre los ecosistemas más amenazados del planeta. La identificación de áreas prioritarias para el establecimiento de Proyectos Integrales de Conservación y Desarrolla (PICD) ha sido restringida debido a la falta de información sobre la distribución, abundancia y requerimientos de hábitat de las especies amenazadas, además del tamaño, condición y amenazas para la supervivencia del bosque remanente. Evaluamos un muestreo rápido de la fauna y procedimientos de modelado de hábitat diseñados específicamente para generar información para la selección de reservas y su diseño, mediante el uso de Sistemas de Información Geográfica (SIG). El estudio fue llevado a cabo en un remanente grande de bosque primario de monson (94,000 ha), utilizando lemures como grupo indicador. La abundancia de los lemures y variables del microhábitat fueron medidas en 64 sitios estratificados de muestreo, dispersos en toda la región. Estas observaciones fueron incorporadas en el SIG con otras variables ambientales y datos de uso del suelo derivados de mapas e imágenes de satélite. Procedimientos estadísticos fueron aplicados para analizar hábitat de especies, predicciones y mapas de distribución de especies, así como para estimar tamaño poblacional de especies a lo largo del área de estudio. Patrones de agricultura con tumba y quema y perturbaciones antropocéntricas fueron también analizados para la identificación de áreas con bajo riesgo de perturbación cultural. Un mapa de distribución de riesgos se sobrepuso a un mapa de biodiversidad relativa de lemures para identificar áreas prioritarias de conservación con alta biodiversidad y bajo riego de perturbación a corto plazo. La diversidad de los lemures estuvo mas fuertemente influenciada por el clareado de hábitat y perturbación humana (cosechas, pastoreo y caza) dentro de una distancia de 8 km de las villas. La diversidad de los lemures fue mas alta en una región con mayor elevación y lluvia, distante de villas y carreteras. La única reserva existente en el área de estudio se localiza en una región con alto riesgo de perturbación. Casi la mitad de la reserva ha sido afectada durante los últimos 45 años. Existe un considerable margen para la reasignación de uso del suelo dentro del área de estudio que incluya un área protegida, una zona de amortiguamiento con caza y cosecha de madera controlada y una instalación adyacente para ecoturismo. Encontramos que la información generada por biodiversidad estratificada y análisis de uso del suelo con una intensidad de muestreo menor al 0.1% es suficiente para proveer la fundación objetiva de planes para biodiversidad regional.
Regurgitation and reingestion (R/R) is a common phenomenon in captive lowland gorillas (Gorilla gorilla gorilla), but it has never been reported in wild populations. Although changes in diet have been shown to reduce the behavior, little is known about the underlying mechanisms that lead to its development and maintenance. In this paper, a comparative analysis of ruminatory behavior in humans and stereotypic behavior in captive, domesticated animals reveals several nutritional and motivational factors that may contribute to the sustained performance of this behavior in captive gorillas. Additional research is needed to evaluate temporal, seasonal, gustatory, social, individual, and medical influences on this behavior.
To study the influence of ‘visitor presence’ on captive lion-tailed macaques, a behavioural study (Short Term Effects) was conducted on 30 individuals housed in 8 zoos across India. The study animals were observed on ‘visitor presence’ days and on ‘visitor absence’ days, which were zoo holidays. To record the long-term impact of ‘visitor presence’ on captive primate behaviour, a study on the Long Term Effects was also conducted. In this study, the behaviour of seven singly-housed individuals was recorded independently when ‘on-exhibit’ and ‘off-exhibit’ in enclosures of similar sizes. In the Short Term Effects, visitors influenced 20% more abnormal behaviour and 3% more social, mating and aggressive behaviours. In the study on Long Term Effects, visitors were found to influence a 30% increase in abnormal behaviour. In addition, in this study on Long Term Effects, use of space in the enclosure varied with individuals using the areas of the enclosure with trees, shrubs and water bodies (enriched zone) more when ‘off-exhibit’ than when ‘on-exhibit’. ‘Visitor presence’ was thus found to influence the behaviour of captive lion-tailed macaques in a negative way suggesting that ‘visitor presence’ might adversely affect their welfare.
1. Relations between sexually mature males in multi-male groups of macaques have been characterised as largely antagonistic, with bonnet macaques being the only frequently cited exception. A survey of the literature indicated that affiliative behaviour between males is more widespread than has been supposed. 2. Factors were examined which may influence the nature of relations between adult males. In all macaque species for which there are adequate data, males commonly move between groups, making it unlikely that kinship between adult males is an important factor in natural groups. 3. Affiliative behaviour between adult males is more frequently observed in small groups, with sex ratios closer to parity, than in large groups with more uneven sex ratios. Two factors are thought to influence this association. First, most large groups with highly skewed sex ratios were provisioned. Provisioning results in increased levels of aggression and tension, which may preclude the formation of affiliative relationships. Second, in groups with highly skewed sex ratios males can find many potential grooming partners among the females and their offspring. Where the sex ratio is more even there may be a shortage of potential partners among the natal animals, and males may form relationships with other males as an alternative. 4. Much of our knowledge of macaque societies has come from provisioned populations. Findings made under such conditions muse be verified with data from natural populations, wherever possible, especially where competition may have an influence on the patterns of behaviours observed.
On top of the scenic escarpment road of A1 Hada, people regularly feed baboons and members of a large troop of Papio hamadryas developed commensal habits, waiting for handouts beside the road. Alarmed by the increasing baboon number, local authorities wish to control the troop’s enlargement. A direct solution to prevent the troop’s growth would be to forbid people to feed baboons. However, both the sufficiency of natural vegetation and the ability of the troop to return to feeding on natural resources is uncertain. The response of baboons to a reduced provisioning was studied when the road was closed for maintenance during winter. We compared the relative share of natural vegetation and handouts in the troop’s diet along the year and monitored range use in relation to food availability. The main effects of handout reduction were the enlargement of daily range combined with relocation and diversification of sleeping sites, the troop splitting into small sleeping groups. Baboons switched easily to a natural diet. Numerous plant species were consumed, but the Juniper species typical of these elevations were the most commonly selected. The results show the feasability and efficiency of the reduction of food provision in managing such commensal groups living partially from human handouts.
Two translocations of commensal rhesus monkey (Macaca mulatta) subgroups were undertaken in India as a conservation measure. In August 1989, a total of 21 animals were transported from Tughlaqabad, a site located on the southeastern outskirts of New Delhi, to Meetha Pur, a rural area about 10 km distant. In February 1990, 13 rhesus were taken from Phari Dheeraj, a crowded residential area in Old Delhi, and released at Lai Kuan, a public parkland located near the border of the State of Haryana. On a per-animal basis, the cost of translocating these 34 monkeys was 70 % less than that realized during a previous relocation of Indian rhesus conducted in 1984. The rhesus is not endangered in India, but does occupy a vulnerable position given that(l) its numbers are much reduced relative to what they were 30 years ago, and (2) it is strongly commensal and capable of causing considerable damage to crops and other property. The utility of translocation in the conservation of India’s remaining rhesus is discussed in the light of the results of the Tughlaqabad and Phari Dheeraj efforts. It is concluded that, while the cost per animal could be cut further by practicing the relocation of commensal rhesus on a larger scale, the use of translocation on a nation-wide basis will probably remain prohibitively expensive in India, a country with limited financial resources.
The activity budget of green monkeys Cercopithecus aethiops was studied in the Guinea Savanna Forest of Old Oyo National Park, Nigeria for 18 months, during both the dry and wet seasons. A focal troop was randomly chosen from the four troops which are resident in the park. The activities studied included travelling, feeding, resting, drinking and grooming. Green monkeys were more active in the morning and evenings than in the afternoons. They travelled and drank more during the dry season than during the wet season, while the least time was spent on grooming during both seasons. Food availability influenced green monkeys' daily activity during all seasons.
Predation pressure has long been proposed as a determinant of mammalian social systems. Group size and composition were compared for 121 populations of cercopithecoid primates, from 39 species of 13 genera, living under low, moderate or high predation risk. In confirmation of previous studies, predation risk was found to have a major effect on group size, with populations under high predation pressure living in significantly larger groups than those at lower risk. However, there were differences in social structure between the predation risk categories. Unimale groups were smaller, had fewer females than multimale groups and were infrequently found under conditions of high predation risk. Predation risk had marked effects on the composition of multimale groups. Under conditions of high predation risk there was a disproportionate increase in the number of adult males over that predicted by the number of females present or by group size. At low predation risk there were fewer males than predicted on the basis of female group size. Together, these results suggest that male strategies to monopolize females depend both on the females' grouping patterns and on the needs of both sexes to maximize group size under high predation pressure.
Among the variety of commensal relationships between human and nonhuman primates, those of most concern today center around « conflict ». As part of a long-term study of olive baboons (Papio anubis) in Kenya, we had the opportunity to investigate the response of four wild troops to the incursion of agriculture. We also tested a variety of management techniques. The development of crop-raiding was not inevitable. In fact, baboons appeared very reluctant to raid, except for a few who had previously fed on human food. Responses ranged from enlarging home range size, decreasing troop size, shifting home range, to raiding as a secondary strategy and raiding as the main foraging orientation. Crops provided greater digestibility and increased foraging efficiency over natural foods. Eating human food altered activity budgets, increased growth and reproduction, but took its toll in injuries and deaths. Control techniques work through increasing costs and decreasing benefits of raiding. The most promising technique is taste aversion conditioning, although better emetics are needed before this method can be widely used. Shifting cultivars may also be effective, particularly if the new crops are less digestible or less palatable for baboons. The prospects for managing primate pests will vary depending on whether animals have viable alternatives. The future of existing primate commensalism is threatened by increasing human pressure on resources. It is highly unlikely that new commensal relationships will develop given these same constraints. Nevertheless, the framework of primate commensalism may provide insights into the variety of human and nonhuman primate relationships and how these might be manipulated for future primate conservation.
Etude conduite le long des rivieres Skagit et Nooksack (Washington). Le canotage perturbe l'approvisionnement des Haliaeetus et provoque des depenses energetiques supplementaires. Mesures de protection a prendre
Four populations of red howling monkeys (Alouatta seniculus) in different states of population growth were studied to examine how changes in intrasexual competition and kin availability influence mating patterns and population genetic structure. The demography of each population was monitored for 5 years. Genetic variance within and between groups and coefficients of relatedness ((r) over bar) were estimated from allozyme loci. Advantages of cooperating with kin over non-kin changed with different demographic circumstances for each sex, such that social groups were more likely to be kin groups when population density was approaching or at carrying capacity than when it was low. In populations growing rapidly through new social group formation, genetic differentiation between groups was low. Genetic differentiation increased as populations approached carrying capacity and groups matured socially (i.e., (r) over bar increased). That trend was reversed as (r) over bar within group breeding subsets (males and females, respectively) increased, thereby decreasing (r) over bar between males and females and reducing group (r) over bar. This was enhanced when inbreeding within groups was followed by transfer of pods of related males, which occurred in a saturated habitat and in a declining population experiencing high mortality of adult females. Heterozygosity within groups in excess of predicted values was increased due to Lineage effects as groups matured and as (r) over bar within breeding subsets increased. It also increased when population growth occurred through immigration from genetically differentiated demes, and when groups fused during population decline. There were large intergroup differences in mortality during population decline, which was accompanied by social instability that resulted in increased intergroup transfer of both sexes, thereby resorting survivors throughout remaining breeding groups. Conditions favoring intergroup selection thus decreased during periods of population growth when selection was relaxed and group (r) over bar was low, increased during periods of stasis when habitats were saturated and group (r) over bar was highest, and were most prominent during periods of population decline when differences in group fitness should have the most pronounced consequences.
The effects of artificial feeding by humans on the behaviour of Hamadryas Baboons (Papio hamadryas) at Al Hada mountain in the western region of Saudi Arabia were examined. Grooming was the dominant behaviour in these baboons, especially in adults throughout all seasons. Play was the dominant activity of juveniles less than 4 years of age. Provisioning the locations changed all categories of behaviour. Less social (in some cases, agonistic) and mounting behaviour were found outside rather than inside provisioning sites. Provisioning periods also changed all kinds of behaviour. Periods of heavy provisioning were characterised by less social behaviour, more play in juveniles and increased agonistic and mounting behaviour in adults.
Kenya is one of the leading tourist destinations in Africa. Most of the country's tourism is based on nature attractions. About 10% of the country has been set aside for wildlife conservation and the promotion of nature-based tourism. This study gives a historical evaluation of western environmental values and how these values influence wildlife conservation and the development of nature-based tourism in Kenya. Also, a comparative analysis is conducted between western environmental values and rural peasants' environmental perceptions.
The impact of road mortality on local
populations of wildlife has rarely been quantified. In June 1991, the access
road into the northern end of the Cradle Mountain – Lake St Clair
National Park in Tasmania was widened and sealed. This occurred part-way
through an ecological study of the dasyurid carnivore guild, during which
populations were being monitored. In 17 months, the resident population of 19
eastern quolls became extinct and the devil population, of 39 individuals,
halved. Concurrently, there was a dramatic increase in the number of
road-kills. The main causal factor was probably an increase in modal speed of
about 20 km h–1 and a greater increase in maximum
speed. Measures were implemented to reduce the incidence of
vehicle/wildlife collisions. Measures directed at people included
physically slowing traffic speed (using ‘slow points’) and
increasing driver awareness (signs and pamphlets). Those directed at wildlife
included deterring wildlife from crossing the road in the path of approaching
vehicles (wildlife reflectors), and encouraging escape off the road (ramps
across gutters and banks, and pipes for shelter). The ‘slow
points’ were effective in reducing vehicle speeds by 20 km
h–1. Wildlife used the ramps and pipes. The
eastern quoll population was re-establishing within six months, and after two
years, had recovered to 50% of its former level. There was some
indication that devil populations were recovering.
For the situation in which the contrasts of interest are limited to the pairwise comparisons among the means of K samples of equal or unequal sizes, four normal univariate single-stage multiple comparison procedures are compared for significance levels not exceeding 0.05: Scheffé's S-method, Dunn's (1, 2] and šidák's [17] improved version of the Bonferroni method, Hochberg's GT2 procedure [8] utilizing the maximum modulus, and Spjøvoll and Stoline's T′-method [19]. Rules are given for determining if any method is uniformly preferable (best for all contrasts). Nonuniform preference rules are also proposed and applied to some examples. Auxiliary tables are provided for selecting a method for significance levels 0.01 and 0.05 for several values of v, the number of degrees of freedom of an independent variance estimate, and K. It is shown that the T′-method is uniformly preferable when the sample sizes are “nearly” equal, while one of the other methods will be uniformly preferable when all sample sizes are “sufficiently” different. The comparison is readily extended to the case in which only a subset of the pairwise contrasts is of interest.
Linnaeus' original scientific description of Lemur catta, the ringtailed lemur, was based on a living animal brought to England in 1749. Although there were many brief descriptions of wild ringtailed lemurs, it was not until Jolly wrote her now classic book, Lemur Behavior, that we had our first detailed description of the natural history of these beautiful animals (Fig. 1). Since then, long-term field studies, mainly from two study sites in Madagascar, Berenty and Beza Mahafaly (Fig. 2), as well as studies on forest-living groups in captivity at the Duke University Primate Center in Durham, North Carolina, have greatly expanded our knowledge of the ecology and behavior of this species (Table 1, Box 1). Thirty-five years of research on this species at these various sites indicates that Lemur catta is proving to be every bit as complex in its behavior as are many anthropoid primates. This very complexity has been reflected in the current controversies and questions concerning the ecology and behavior of this species.
Habituation typically is viewed as a negative consequence of human interactions with wildlife (Higginbottom, 2004; Newsome, Dowling, & Moore, 2005; Shackley, 1996). While animal habituation commonly is used in the laboratory and field-based zoology studies, attempts to consider deliberate habituation specifically in a tourism management context (Shelton, Higham, & Seddon, 2004) has been received unsympathetically by biological scientists and wildlife managers on the grounds that habituation, by definition, is undesirable. This paper puts forward the case that the global and stable behavioural descriptor, habituation, is not the most useful way to formulate most observed lack-of-wildlife-response to visitor approach and observation. It presents an applied behaviour analysis of wildlife habituation that is situated within learning theory. This analysis differentiates between avoidance/approach behaviours, tolerance, habituation and sensitisation. This provides a formulative framework for human–wildlife interactions, that is then considered specifically in terms of tourism businesses seeking to provide sustainable visitor interactions with wild animals. A tourism management model derived from this critique of habituation is presented and discussed.
There are 2 main competing theories on the evolution of group living in diurnal non-human primates. 1) Predation avoidance favours group living; there are only disadvantages to feeding in a group and feeding competition increases with group size. 2) There is a feeding advantage to group living deriving from communal defence of high-quality food patches; predation is not important. A critical test is proposed: the theories differ in the predicted relationship between a female's birth rate and the size of the group in which she lives. An additional test is concerned with the predicted relationship between population density relative to food availability and average group size. Finally, a critical test is proposed of the hypothesis that increasing group size should lead to reduced predation risk by comparing demographic patterns between areas where predators are still present and where they have disappeared. In all 3 tests, results provide strong support for the predation-feeding competition theory and are clearly unfavourable for the theory postulating feeding advantages to group living. Such feeding advantages may, however, gain prominence under some conditions.-from Author
This research examines the relationship between visitor numbers and the behaviour of Diana monkeys (Cercopithecus diana diana). A family group of two adult and four juvenile captive Diana monkeys was studied intensively for 10 days at Edinburgh Zoo, UK. The six subjects were scan sampled every 5min for 7h per day; their behaviour, and visitor numbers, were recorded. Chi-squared analysis revealed a highly significant association between Diana monkey behaviour and visitor group size, where the amount of time spent grooming and sleeping/resting decreased with greater visitor numbers whereas time spent playing and feeding/chewing increased. Results are discussed in the context of recent findings regarding visitor presence and zoo animal welfare.
Activity budgets and social aspects of feeding, among the adult females in a group of semiprovisioned baboons that fed from a garbage dump were compared with those of adjacent wild-feeding groups in Amboseli National Park, Kenya, during the dry seasons of 1984 and 1985. Statistically significant differences were found in time spent feeding, distance travelled, and the relationship between dominance rank and time spent feeding. The garbage-feeding animals fed for 20% of the time and rested for almost 50%, in contrast to approximately 60% and 10%, respectively, for the wild-feeding animals. Speed of travel, length of day-route, and home range size were greatly reduced for the garbage-feeding animals. Use of sleeping trees and day route were highly regular in contrast to the unprovisioned group. At the garbage dump, time spent feeding was correlated with dominance rank among the adult females of this study. This was not the case for feeding on wild foods. Human enriched food sources offer the opportunity to study limiting factors and relationships between ecology and behavior. However, these conditions lead to human–animal conflicts that may be to the animals' long-term detriment. Conservation and management implications are discussed.
Although the control mechanism is less clearly determined, stress in birds, as in mammals, can be defined in terms of the activation of two integrative neurohumoral systems by external or internal stimuli: (a) the neurogenic system response, which is almost immediate and provides a rapid delivery of energy, and (b) the hypothalamic-pituitary-adrenal complex response, which is a slower, long-term mobilization of defenses. These responses, although temporary defense mechanisms against specific stimuli, place a bird in a state of general nonspecific stress, in which growth rates and resistance to many diseases are diminished.
This study monitored the impact of habituation and tourism on a community of chimpanzees in the Kibale Forest, Uganda. From February 1992 to June 1993, chimpanzees' initial reactions to contacts with human observers were recorded together with other factors such as numbers of observers present and the method used to locate the chimpanzees. On 54 occasions with various numbers of observers (and/or tourists) present, activity rates were recorded for the observed individuals. Numbers of observers present (up to five) had no impact on the chimpanzees' initial reaction, and the only significant change in activity caused by higher numbers of observers (up to 15) was an increasing vocalisation rate. As visitor satisfaction declines with groups of more than six to eight people it is suggested that tourist groups viewing chimpanzees should be kept at a maximum number of 10 (including guides).
Behavioral and adrenocortical responses of rhesus (Macaca mulatta), bonnet (M. radiata), and crabeating (M. fascicularis) macaques were compared in their home cages, during exposure to novelty and during physical restraint. Both behavioral and adrenocortical responses differentiated species in each condition. In all conditions, post-test corticosteroid levels were highest for crabeaters and lowest for rhesus. Rhesus were the most active behaviorally, and bonnets were the most passive, while crabeaters exhibited the greatest signs of behavioral disturbance. Relationships between adrenocortical and behavioral responses varied between groups. Both adrenocortical and be- havioral profiles were in accord with the behavior of these three species under more natural conditions. The role of psychophysiological responses in general behavioral dispositions toward the environment is discussed. It is concluded that behavioral dispositions, inclusive of psychophysiological re- sponses, may vary qualitatively even among closely related primate species.
Animals can perceive tourists as predators and will incur fitness costs should their predator-avoidance behaviours result in forgone resource acquisition. Not all wildlife, however, treat tourists as predators; animals can respond positively to tourists, especially when food is used as an attractant. We investigate the costs posed by novel grouping over a tourism-provisioned food resource in a normally solitary forager, the southern stingray Dasyatis americana, in Grand Cayman. Specifically, we test the hypotheses that group-living stingrays in a new environment — which includes both the presence of tourists and quickly renewing food patches — will be exposed to increased injuries, ecto-dermal parasites and aggressive interference competition that result from the unusual grouping behaviour. We found that, in comparison to stingrays from non-tourist sites, tourist- fed stingrays are more likely to have lower body condition, be injured by boats and predators, be sus- ceptible to ecto-dermal parasites, and be engaged in intense interference competition (in the form of conspecific bite marks). Stingrays from tourist sites also have significantly higher mean numbers of injuries, parasites, and median bite marks. By exploring alternative hypotheses to explain the pattern of our findings (e.g. natural causes/behaviour), we unequivocally show that the impacts incurred by the stingrays stem from the effects of tourism. These findings suggest that novel grouping poses costs to the stingrays; the tourist site represents a riskier habitat with regards to injury and predation; and there may be long-term fitness consequences. From a management perspective, measures should be taken to alleviate the crowded conditions at tourist sites, in terms of both boat and stingray density.
Visitors to zoos can be a source of potential stress to some captive-housed primates. To date, the effect of the human audience on the welfare of captive gorillas has been overlooked. This study thus examined the behaviour of six zoo-housed gorillas during periods of both high and low visitor density, to establish whether there was any effect of audience numbers on the animals’ well-being. Low visitor density encouraged behaviour suggestive of relaxation, with gorillas spending a significantly greater proportion of time resting. High visitor density, by contrast, encouraged significantly more intragroup aggression, stereotypies and autogrooming. The findings suggest that gorillas, like many other species of primate, are excited by high numbers of visitors.
Data were collected on the behavior and physical characteristics of 259 human visitors and 24 chimpanzees at Chester Zoo. The successive responses of humans and chimpanzees to each other's behavior were recorded, the resulting long sequence being referred to as an interaction sequence. There was no particular set of characteristics that distinguished interactors from noninteractors in either humans or chimpanzees, although there was some evidence that chimpanzees were particularly likely to respond to men carrying objects. Chimpanzee responses were random with respect to the previous human behavior, but human responses were significantly associated with the preceding chimpanzee behavior. In particular, chimpanzee sounds were likely to be followed by human sounds, and begging was likely to be followed by the offer of food. Interaction sequences varied in length, but 9% of chimpanzee-initiated sequences went as far as a ninth interaction. Sequences resulted in the chimpanzees being given food in 25% of human-initiated, but only 8% of chimpanzee-initiated sequences. The results are consistent with the interpretation that humans and chimpanzees are motivated to interact with one another and that the chimpanzees do this primarily to obtain food. © 1995 Wiley-Liss, Inc.
Fraser Island is a unique tourist destination catering to more than 300,000 visitors annually. A significant attraction of the World Heritage-listed island are the dingoes, which are regarded as one of the purest strains in Australia. The dingo population is estimated to be between 150 and 200 animals and their conservation is of national significance. Concerns have long been expressed about the potential for dangerous interactions between dingoes and humans. As visitation to the Island has increased, aspects of the environment have changed and dingoes have altered their normal habits. Management practices have focussed on removing unnatural food sources, such as open rubbish dumps, and educating the public. On April 30, 2001, dingoes mauled a 9-year-old boy to death and the public demanded firm management decisions. However, the fundamental question remained. Do we manage the animals or the people? Public opinion was polarised. This article describes the issues and politics of managing dingoes and tourists on Fraser Island, and highlights how a single serious incident can influence management decisions.
During the last two decades, much research has focused on the mechanisms used by nonhuman primates for conflict resolution. Reconciliation, i.e., a friendly reunion between former opponents, has been reported in several primate species. Reconciliation seems to serve at least two functions. According to the Valuable Relationship Hypothesis, reconciliation restores the disturbed relationship between former opponents and, consequently, occurs more often between individuals with more valuable relationships. The Uncertainty-Reduction Hypothesis emphasizes the function of reconciliation to reduce anxiety and uncertainty in the recipient of aggression following a conflict. The study of post-conflict emotionality facilitates the integration of these two hypotheses. The present study focuses on the factors affecting post-conflict anxiety of the two former opponents. Post-conflict data of captive Barbary (Macaca sylvanus) and long-tailed macaques (Macaca fascicularis) are used here to examine patterns of anxiety-related behavior, such as self-scratching. The fact that in the first minutes following a conflict not only recipients of aggression but aggressors too increased the rate of scratching above baseline levels suggests that the risk of renewed attacks is unlikely to be the only reason for post-conflict anxiety. The quality of the relationship between former opponents was a good predictor of post-conflict scratching rates by the recipient of aggression: Rates were higher after conflicts between individuals with stronger affiliative relationships. This result was not due to differential risk of resuming hostilities or arousal resulting from the intensity of the previous conflict. This finding suggests that the disturbance of a valuable relationship due to the previous conflict (e.g., the potential loss of benefits provided by such relationships) is a major cause of post-conflict anxiety. In line with the Uncertainty-Reduction Hypothesis, the best way to cope with post-conflict anxiety is to reconcile and therefore restore the relationship. Higher levels of post-conflict anxiety should lead therefore to higher conciliatory tendency. This interpretation is supported by the fact that individuals with strong affiliative relationships reconcile more frequently than others, as predicted by the Valuable Relationship Hypothesis. The mediating role of emotions, such as post-conflict anxiety, is probably not limited to reconciliation. Other post-conflict interactions, such as consolation, triadic reconciliation, and mediation, are also likely to be better understood if future studies focus on the post-conflict emotions of the opponents and bystanders. Aggr. Behav. 23:315–328, 1997. © 1997 Wiley-Liss, Inc.
Simulations are necessary to assess the performance of home-range estimators because the true distribution of empirical data is unknown, but we must question whether that performance applies to empirical data. Some studies have used empirically based simulations, randomly selecting subsets of data to evaluate estimator performance, but animals do not move randomly within a home range. We created an empirically based simulation using a behavioral model, generated a probability distribution from those data, and randomly selected locations from that distribution in a chronological sequence as the simulated individual moved through its home range. Thus, we examined the influence of temporal patterns of space use and determined the effects of smoothing, number of locations, and autocorrelation on kernel estimates. Additionally, home-range estimators were designed to evaluate species that use space with few restrictions, traveling almost anywhere on the landscape. Many species, however, confine their movements to a geographical feature that conforms to a relatively linear pattern. Consequently, conventional analysis techniques may overestimate home ranges. We used simulations based upon coastal river otters (Lontra canadensis), a species that primarily uses the aquatic-terrestrial interface, to evaluate the efficacy of fixed and adaptive kernel estimates with various smoothing parameters. Measures of shoreline length within contours from fixed kernel analyses and the reference smoothing parameter were best for estimates of 95% home ranges, because smoothing with least squares cross validation (LSCV) often resulted in inconsistent results, excessive fragmentation, and marked underestimates of linear home ranges. Core areas (50% density contours) were best defined with fixed kernel LSCV estimates. Fewer locations underestimated linear home ranges, and there was a subtle positive relation between home-range size and autocorrelation. Generally, as location numbers increased, autocorrelation increased, but differences from the "true" home range decreased. Results were similar for our simulations and empirical data from 13 river otters. Examination of empirical data revealed that data with high positive autocorrelation illustrated seasonal reproductive activities. Because autocorrelation does not negatively influence estimates of linear home ranges, assessment of independence between data points may be more appropriately viewed as a means to identify important behavioral information, rather than as a hindrance.
Seven major types of sampling for observational studies of social behavior have been found in the literature. These methods differ considerably in their suitability for providing unbiased data of various kinds. Below is a summary of the major recommended uses of each technique: In this paper, I have tried to point out the major strengths and weaknesses of each sampling method. Some methods are intrinsically biased with respect to many variables, others to fewer. In choosing a sampling method the main question is whether the procedure results in a biased sample of the variables under study. A method can produce a biased sample directly, as a result of intrinsic bias with respect to a study variable, or secondarily due to some degree of dependence (correlation) between the study variable and a directly-biased variable. In order to choose a sampling technique, the observer needs to consider carefully the characteristics of behavior and social interactions that are relevant to the study population and the research questions at hand. In most studies one will not have adequate empirical knowledge of the dependencies between relevant variables. Under the circumstances, the observer should avoid intrinsic biases to whatever extent possible, in particular those that direcly affect the variables under study. Finally, it will often be possible to use more than one sampling method in a study. Such samples can be taken successively or, under favorable conditions, even concurrently. For example, we have found it possible to take Instantaneous Samples of the identities and distances of nearest neighbors of a focal individual at five or ten minute intervals during Focal-Animal (behavior) Samples on that individual. Often during Focal-Animal Sampling one can also record All Occurrences of Some Behaviors, for the whole social group, for categories of conspicuous behavior, such as predation, intergroup contact, drinking, and so on. The extent to which concurrent multiple sampling is feasible will depend very much on the behavior categories and rate of occurrence, the observational conditions, etc. Where feasible, such multiple sampling can greatly aid in the efficient use of research time.
There is growing interest in examining the effects that visitors have on the welfare of zoo primates. Most recently, the importance of evaluating the combined impacts of visitors and management practices on anxiety levels of these animals has been highlighted. Studies that take such an approach have the potential not only to highlight how visitors affect the welfare of zoo primates, but also how these impacts may be mediated by management interventions, such as enrichment. The aim of this study was to explore the potential impact of visitor numbers on behavioural indices of anxiety among western lowland gorillas in two UK zoos, Port Lympne and Chessington, and to determine whether feeding enrichment mediates any such visitor effects. During fifteen minute focal watches, visitor numbers were assessed and all occurrence data collected on two behavioural indices of anxiety: self-scratching and visual monitoring of visitors. Data collected at each site were divided according to whether they were taken during periods of feeding enrichment, or during periods when no such enrichment was being given. Analyses revealed no evidence for a visitor effect at Chessington, with durations of self-scratching and visual monitoring unrelated to visitor number, either during periods of enrichment or outside of such times. At Port Lympne, durations of both self-scratching and visual monitoring were positively associated with visitor number when no feeding enrichment was taking place; no such relationships were seen, however, during periods of feeding enrichment. Potential reasons for the difference between sites in the occurrence of visitor effects, and for the absence of visitor effects during feeding enrichment at Port Lympne are discussed. The results of this study highlight the importance of assessing visitor effects on the same species at multiple sites, and also suggest that such effects may be reduced by positive management approaches.
The quantification of behaviours linked to anxiety or stress provides a powerful means to address applied questions related to the well-being of captive animals. This study explored correlates of two such behaviours – self-directed (SDB) and stereotypic behaviour (STB) – among captive red-capped mangabeys, Cercocebus torquatus torquatus. Study animals were held at two sites run by CERCOPAN, a primate sanctuary in Nigeria. At the first site (Calabar), animals were housed in traditional cage enclosures, while at the second (Rhoko), they lived in a semi-free ranging environment. Analyses revealed that while animals at the two sites did not differ in time spent in SDB (self-scratching, self-grooming, yawning and body shaking), animals at Calabar showed a higher prevalence of, and spent more time in STB (pacing and head rolling). There were no significant differences in time spent in SDB or STB between the sexes, or between captive-born and wild-born animals. A positive correlation was found between age and time spent in SDB, but not between age and time spent in STB. Finally, positive relationships were found between time spent in SDB and both time spent grooming and time spent being groomed; no relationships were found between grooming behaviour and STB. As stress and anxiety can have detrimental effects on psychological well-being, physical health and reproduction, these findings have important implications for the captive welfare and conservation of red-capped mangabeys, and also potentially of other endangered vertebrate species.