The effects of food sources on Japanese monkey home range size and location, and population dynamics

The effects of supplemental feeding by tourists on wild Japanese monkey's home range size and location, and troop size and composition were studied for two monkey troops, Troop A and Troop B, living along the Irohazaka loop road, Nikko National Park, central Japan. Changes were documented based on data gathered from 1982 to 1996 by the use of radio telemetry. Troop A's home range size shrank and changed from separate winter and summer ranges to a single, year-round home range, with its core located in a high elevation area where supplemental feeding by tourists was heavy. Troop B's home range also shrank and shifted to a lower elevation where supplemental feeding by tourists was heavy. Troop A's population size increased between the winters of 1983–1984 and 1990–1991 in conjunction with an increase in human encounter rates, and then decreased. Troop B's size increased until the winter of 1993–1994, and then decreased. The instability of troop size between 1993 and 1996 may be explained by documented factors such as a decrease in the adult sex ratio, an increase in the infant-female ratio, and an increase in juvenile mortality and/or emigration, all of which may have been influenced by supplemental feeding by tourists.     

Scent marking as resource defense by female Lemur catta

Because ring-tailed lemurs (Lemur catta) are a female-dominant, female-philopatric species in which the females provide the majority of parental care and troop defense, resource defense is a possible function of female lemur scent marking. To test this hypothesis, I conducted three studies. First, I presented captive, individually housed females with a series of samples of female scent, each from a different female, to determine whether they would respond to those samples and discriminate between them. Second, I reanalyzed data from a focal animal study of four females in two adjacent troops in Berenty Reserve, Madagascar, to determine female marking rates before, during, and after the mating season, and to clarify the relationship among positions of feeding, intertroop defense, and scent marking. The third study was based on ad libitum observations of the sniffing and marking behavior of a troop in Berenty Reserve during a year when they traveled far out of their home range. The females in study 1 investigated female scent samples but provided no evidence that they discriminated between them. In study 2 the wild females marked throughout the study and did not limit their marking to the mating season. They deposited significantly more of their marks in a zone of confrontation with adjacent troops, where they also did the majority of their feeding, and they increased their rate of marking during agonistic intertroop confrontations. The females determined the positions of their scent marks and deposited the first mark in the majority of countermarking sequences. When the females traveled out of their defended range in study 3, they significantly decreased their rate of marking and increased their rate of sniffing spots but not marking them. All evidence gathered so far supports the hypothesis that one function of female ring-tailed lemur scent marking is to provide intergroup information that is then used to reinforce the border of the defended resource.     

Meadow Voles (Microtus pennsylvanicus) and Prairie Voles (M. ochrogaster) Differ in Their Responses to Over-Marks from Opposite- and Same-Sex Conspecifics

Over‐marking occurs when one individual deposits its scent mark on the scent mark of a conspecific. Previous studies have shown that meadow voles (Microtus pennsylvanicus) and prairie voles (M. ochrogaster) that were exposed to an over‐mark of two same‐sex conspecifics, later responded similarly to the top‐scent mark but differed in their response to the bottom‐scent mark. In the present study, we examined the responses of meadow voles and prairie voles to same‐sex and mixed‐sex over‐marks to ascertain whether their responses reflect the different tactics which males and females in promiscuous (meadow voles) and monogamous (prairie voles) species use to attract opposite‐sex conspecifics and to compete with same‐sex conspecifics. Males and females of both species spent more time investigating the mark of the top‐scent donor than that of the bottom‐scent donor of an over‐mark. Meadow voles exposed to a mixed‐sex over‐mark spent more time investigating the mark of the opposite‐sex conspecific independently of whether it was from the top‐ or bottom‐scent donor. In contrast, prairie voles spent more time investigating the mark of the opposite‐sex donor if it was from the top‐scent donor. These results suggest that: (i) over‐marking serves a competitive function; (ii) the scent marks of individuals attract multiple mates in promiscuous species such as the meadow vole; and (iii) the scent marks of individuals establish and maintain pair bonds between familiar opposite‐sex conspecifics in monogamous species such as the prairie vole.     

Observations onCebus capucinus in Southwestern Panama

Observations onCebus capucinus in 10 forests in Chiriqui, Panama, are reported. Troop size ranged from two to five in small forests to 20 or more in large forests. A 10 week study was conducted in a coastal forest that contained a troop of 27 to 30Cebus. The troop’s daily path length was approximately 1.6 to 2.5 km and their home range was between 32 and 40 ha. TheCebus usually spent 50 to 70% of their day foraging and traveling. On one occasion, aCebus attacked anIguana, wrestled with it, eventually broke off 30 to 40 cm of its tail, and ate meat from the tail. The relations betweenCebus andSaimiri oerstedii, Alouatta palliata, and man are described.     

Meadow Voles (Microtus pennsylvanicus, Arvicolidae) Over-mark and Adjacent-mark the Scent Marks of Same-sex Conspecifics

Scent over-marking occurs when an animal deposits its scent mark on top of the scent mark of a conspecific; adjacent-marking occurs when an animal deposits its scent mark next to the scent mark of a conspecific. Given that male rodents usually scent mark more than females and that animals spend more time investigating the odor of the top-scent donor of an over-mark, I tested the following three hypotheses. First, male meadow voles deposit more scent marks than female meadow voles. Second, male meadow voles will deposit more over-marks and adjacent-marks in response to the scent marks of a same-sex conspecific than females would. Third, meadow voles spend more time investigating the odor of the second vole placed in the arena than that of the first vole placed in the arena. To test these hypotheses, two age-matched, like-sex conspecifics (first vole and second vole) were placed successively into an arena in which they were allowed to freely explore and scent mark for 15 min. The first hypothesis was not supported. The first and second vole, independently of sex, deposited a similar number of scent marks. The second hypothesis was also not supported by the data: more conspecific scent marks were over-marked by the second female than by the second male. The third hypothesis was supported by the data. After investigating a scented arena, males and females spent more time investigating the odor of the second vole than that of the first vole. Sex differences in scent-marking behaviors of meadow voles are unlike those reported for other species of rodents.     

Dietary protein content affects the response of meadow voles, <Emphasis Type="Italic">Microtus pennsylvanicus</Emphasis>, to over-marks

The response to signals, including scent marks, from opposite-sex conspecifics can be affected by the nutritional state of both the sender and receiver of these signals. Protein content of the diet affects how meadow voles (Microtus pennsylvanicus) respond to single scent marks, but it is unknown how it affects an individual’s response to the overlapping scent marks of two donors (an over-mark). In experiment 1, we tested the hypothesis that protein content of the diet affects the amount of time voles spend investigating the marks of the top- and bottom-scent donors of an over-mark. Males and females fed a 22% protein diet spent more time investigating the scent mark of the top-scent donor than that of the bottom-scent donor; voles fed 9% and 13% protein diets spent similar amounts of time investigating the top- and bottom-scent donors. In experiment 2, we tested the hypothesis that protein content of the diet of the top- and bottom-scent donors affects the amount of time conspecifics spend investigating their scent marks. Female voles spent more time investigating the mark of the top-scent male than that of the bottom-scent male, independent of the differences in protein content of the diets of the top- and bottom-scent donors. Male voles, however, spent more time investigating the top-scent female when she was fed a diet higher in protein content than that of the bottom-scent female. Our results are discussed within the context of the natural history of voles.     

Group composition,home range,and diet of the maroon leaf monkey (Presbytis rubicunda) at Tanjung Puting Reserve,Central Kalimantan,Indonesia

Maroon leaf monkeys (Presbytis rubicunda) were studied at the Orangutan Research Conservation Project study area at the Tanjung Puting Reserve in Central Indonesian Borneo for approximately 15 months in 1974, 1975, and 1977. Over 250 observation hr involving 300 encounters with the monkeys were logged. Troop size ranged from three to eight individuals in the nine troops surveyed with only one adult male being present in each troop. Home range size varied from 0.33 km² to 0.99 km² and correlated with troop size. Forty-six different food types were observed eaten by the maroon leaf monkeys; of these foods, 52% consisted of fruits, 36% of leaves, and 12% of flowers.     

Take-over and infanticide in South Indian Hanuman langurs (Presbytis entellus)

A take-over and infanticide were observed in a South Indian population of Hanuman langurs (Presbytis entellus). A one-male, bisexual troop of langurs (Troop B1) was attacked by a band of two adult males and one subadult male. During the take-over, the resident male and one elderly female were driven from the troop. The youngest infant, about two months old, was fatally wounded by an attack from the dominant incoming male (S1). However, other young infants in Troop B1 were left unharmed. All three incoming males remained in Troop B1 but the troop appeared to remain functionally one-male, as S1 did not allow the other adult male to copulate. There was no sign that S1 attempted to eject the other two males from troop B1. © 1993 Wiley-Liss, Inc.     

Solitary male chacma baboons in a desert canyon

Solitary and paired adult (nine) and subadult (one) male chacma baboons, Papio ursinus, were observed over a period of years living in part of a wooded desert canyon not used by adjacent troops. These extratroop males were silent when alone and gave only one alarm vocalization, the “wa-hoo” call, when paired. The space occupied by them is unsuitable for use by troops according to criteria for adequate sleeping sites and access to water. But the foods available to them, especially figs, but also other fruits and fresh acacia seeds, were abundant. These foods are more highly preferred by baboons than those foods available to troop members. Troop members deplete these resources and shift to less preferred foods with lower water content and longer processing times. All of the adult members of the troop adjacent to these isolated males were infected with a skin disease. Isolated males were not so afflicted and so cannot have originated from, or ever been a part of, this troop. They probably moved to the space where they were observed from other inland troops, traveling to their current home range along the narrow canyon river course.     

Olfactory Experience Affects the Response of Meadow Voles to the Opposite‐Sex Scent Donor of Mixed‐Sex Over‐Marks

Scent marking and over‐marking are important forms of communication between the sexes for many terrestrial mammals. Over the course of three experiments, we determined whether the amount of time individuals investigate the scent marks of opposite‐sex conspecifics is affected by 4 d of olfactory experience with those conspecifics. In Experiment 1, female meadow voles, Microtus pennsylvanicus, spent more time investigating the scent mark of the novel male conspecific than that of the familiar male donor, whereas male voles spent similar amounts of time investigating the scent mark of the familiar female and a novel female conspecific. In Experiment 2, voles were exposed to a mixed‐sex over‐mark in which subjects did not have 4 d of olfactory experience with either the top‐scent donor or the bottom‐scent donor. During the test phase, male and female voles spent more time investigating the scent mark of the opposite‐sex conspecific that provided the top‐scent mark than that of a novel, opposite‐sex conspecific. Male and female voles spent similar amounts of time investigating the scent mark of the bottom‐scent donor and that of a novel opposite‐sex conspecific. In Experiment 3, voles were exposed to a mixed‐sex over‐mark that contained the scent mark of an opposite‐sex conspecific with which they had 4 d of olfactory experience. During the test phase, male voles spent more time investigating the mark of the familiar, top‐scent female than the scent mark of a novel female donor but spent similar amounts of time investigating the mark of the familiar, bottom‐scent female and that of a novel female donor. In contrast, female voles spent more time investigating the mark of a novel male donor than that of either the familiar, top‐scent male or that of the familiar, bottom‐scent male. The sex differences in the responses of voles to scent marks and mixed‐sex over‐marks are discussed in relation to the natural history and non‐monogamous mating system of meadow voles.     

Population changes and ranging behaviour of wild Japanese monkeys at Mt. Kawaradake in Kyushu,Japan

Population changes and home range utilization of the wild Japanese monkey at Mt. Kawaradake have been studied since 1972. Age compositions of this troop were obtained over a seven-year period. Troop size decreased from over 100 to 40 individuals as a result of a capture in 1974. The capture affected directly and indirectly the troop's range and population dynamics. The troop reduced its range size from 4.7 km² to 2.67 km² and changed its utilization pattern in relation to the decrease in size. After the capture, the troop used one particular area intensively, whereas the rhythmic nomadic pattern had been observed as before. This may have been caused by the decrease in the overall food requirement of the troop. The birth rate increased significantly after the capture. However, troop size did not increase because of the low recruitment rate for adult females and the high mortality of juveniles.     

Female preference for males that have exclusively marked or invaded territories depends on male presence and its identity in the subterranean rodent Ctenomys talarum

Territorial scent‐marking provides chemical records of male competitive interactions that are available to females, who gain valuable information to assess and identify best quality partners. In this context, the solitary subterranean rodent tuco‐tuco (Ctenomys talarum) offers excellent possibilities to evaluate the effects of male exclusive scent‐marking of territories on female assessment. For evaluation, we used wild caught individuals of C. talarum, manipulated their scent marks within the territories in captive conditions and staged preference tests where females were able to choose between exclusive and invaded territories. The evaluation was performed in two scenarios considering the identity of the intruder scent mark: territories invaded by a strange male and territories invaded by a neighbour male. Females investigated the chemical cues deposited on the substrate of the exclusively marked territory more frequently. Next, females displayed equal interest to scent samples of both males presented in a Y‐maze. Finally, when females could gain access to both individually isolated males and their scent‐marked territories, they spent more time within invaded territories despite they visited them with the same frequency. Moreover, females tried to get in contact by scratching the mesh of the owner of the invaded territory more frequently. We found that females of C. talarum evaluate the homogeneity (exclusiveness) of scent marks within a male territory and then show preferences in relation to the identity of the intruder's scent –whether strange or neighbour.     

Self-Discrimination in Meadow Voles, Microtus pennsylvanicus

Particular features of the signaling characteristics of the scent marks of temperate zone, seasonally breeding mammals may reflect differences in their reproductive state and, hence, be variable. Consequently, an individual's perception of self may depend more on the condition independent than on the condition-dependent signaling characteristics of the scent marks. Yet, we do not know whether an individual responds to changes in the signaling characteristics of its own scent marks, such as those associated with changes in an individual's reproductive state. Such changes may affect how and where an animal scent marks. Here we report on a series of experiments designed to test the hypothesis that individual meadow voles, Microtus pennsylvanicus , distinguish between scent marks they deposited when they were in different reproductive states. Results showed that voles discriminated their own scent marks from those of unfamiliar, same-sex conspecifics, and the scent marks of siblings. Voles did not behave as if they could distinguish between their own scent marks if the marks were deposited when the voles were in the same reproductive state, although the two scent marks used as stimuli differed in age by 30 d. However, they did so distinguish if they were exposed to scent marks taken when they were in different reproductive states. Overall, these findings suggest that voles behave as if their novel and familiar scent marks shared the similar signaling features. If, however, the reproductive condition of the voles differed when it provided the two scent marks, they behaved as if their own scent marks had different signal characteristics, which may have induced voles to treat the two scent marks as not being the same or having been deposited by two different donors. We speculate that the scent marks of individuals may have unique signaling characteristics that may be associated with that individual's 'current template for self'.     

Selection of scent-marking sites by klipspringers (Oreotragus oreotragus)

Klipspringers deposit scent marks in the form of secretion from the preorbital glands on branches of low trees and shrubs within their territories. Local environmental factors which affect choice of scent-marking sites in a population of wild klipspringers in Zimbabwe are described. Scent marks were typically placed within a narrow vertical range at the end of upward-pointing twigs. Twigs were more likely to be selected if they were located above bare rock, particularly if the ground was raised above the immediately surrounding area or was close to a significant break in slope. A substantial proportion of sites were placed on dead wood. Marking sites may remain in use for a number of years, with turnover rates calculated at between 1 and 2% per annum. The species of tree on which marks were placed were recorded and compared with overall abundance in the habitat. Palatable tree species were found to be selected preferentially when compared to unpalatable ones. Several explanations for this are compared. It is concluded that, simply through preferential selection of certain local characteristics, klipspringers may enhance the visibility and predictability of their scent marks.     

Socio-ecological study ofPresbytis aygula in West Java

Field studies of the Sunda Island leaf monkey,Presbytis aygula, were made in the montane forests of West Java at an altitude of 1,400–1,800 m between September 1976 and August 1981. The troop size ofP. aygula varied from 3 to 12 animals and the population density was about 35 animals/km² in Patenggang and 11–12 animals/km² in Kamojang. They were organized essentially in one male troops, though the troops in the Patenggang area usually consisted of only an adult pair and its offspring. Home range size was about 14 ha in Patenggang and about 35–40 ha in Kamojang. Some of those ranges overlapped with each other. The differences between these two areas were considered to be the result of recent habitat destruction and human impact in Patenggang. Animals spent more time in resting than other activities during a day.P. aygula in Java, consumed mainly leaves, eating less fruits and other parts of plants. Troop cohesion was tight, but social interactions between troop members were rather infrequent. The patterns of intertroop encounters and vocal sounds were described.     

Troop Histories and Range Inertia of Lemur catta at Berenty,Madagascar: A 33-year Perspective

Lemur catta troops in a 1-km ² study area at Berenty Reserve have maintained fidelity to core areas since Budnitz and Dainis' study of 1972–1973, and for two troops possibly since 1963. Population in 1 km ² fluctuated from 155 to 105 to 282 individuals (excluding infants), and the number of troops increased from 12 to 21. Most troops retain the same core areas from year to year (170 observed troop-years). Ten troops derived from known fissions have settled in parts of their parent troop range or an adjacent neighbor's range. Five more troops may derive from similar matrilocal fissioning, inferred from behavior and ranging patterns. One has remained unchanged. Five have unknown parentage, in the ranges of four previously censused troops. Once a fissioned troop completely replaced another, one troop permanently extended its range, three times females joined a different troop, once a female remained nomadic for two years without stable home range. No fissioned troop has been seen to leapfrog others: to settle discontiuously from its parent. Intertroop antagonism may reflect benefits of long-term core area control.     

Troop fission in femaleLemur catta at berenty reserve,Madagascar

A- Troop of Lemur cattaat Berenty Reserve, Madagascar, divided in July- August 1992. We observed the daughter troops— Al and A2— between Aug. 15 and Nov. 15. The subordinate female subgroup of A- Troop (A2) had been harassed for at least 3 years previously. After fission, it suffered high conflict and energy demands, wounding, and socially induced infant mortality. Costs of fission may be a pressure for remaining in large troops. In September, A2 ranged widely, was chased by all neighbor troops, but then it challenged the previously dominant subgroup, Al, and by late October occupied a portion of the previous A- Troop range in which it was consistently successful in confrontations. A2's acquisition of range may relate to its high affiliation and low conflict between females of the subgroup. Unlike previously reported fission in L. cattathere was no male involvement in defense, and the females that lost infants became more aggressive. This may be interpreted as female competition for resource control.     

Over-marking and Adjacent Marking are Influenced by Sibship in Male Prairie Voles,Microtus ochrogaster

Scent over-marking occurs when an animal deposits its scent mark on top of the scent mark of a conspecific. Over-marking may provide advantages in the transfer of information to the individual whose scent is on top but not to the individual whose scent is on the bottom. We tested the hypothesis that over-marking is a competitive form of olfactory communication and that male prairie voles would over-mark the scent marks of same-sex conspecifics more than those of same-sex siblings. Two age-matched male voles (first male and second male) were placed successively into an arena in which they were allowed to explore freely and scent mark for 15 min at age 12, 20, 28, 36, 44, and 52 d. The first male was placed into a clean arena, whereas the second male was placed into an arena containing either the scent marks of an age-matched male sibling or nonsibling. Age affected the total number of scent marks deposited by the voles; 12-20-d-old voles deposited fewer scent marks, over-marks and adjacent marks than did 28-52-d-old voles. Sibship did not affect the total number of scent marks deposited by the first and second voles but did affect the number of over-marks and adjacent marks deposited by the second vole. Siblings received significantly fewer over-marks and adjacent marks than did nonsiblings; this effect was most dramatic after the voles reached 28 d of age, a time coincident with the onset of puberty. Males separated from siblings and housed singly at 44-d-old and tested at 52-d-old, deposited significantly more over-marks and adjacent marks in arenas if the first vole was a nonsibling than if it was a sibling. This differential scent-marking supports the hypothesis that over-marking and adjacent marking are used as competitive forms of olfactory communication by male prairie voles.     

Scent marking and mate choice in the prairie vole, Microtus ochrogaster

Scent marking is common among male and female rodents and might be used in male-male competition and as a mechanism for mate attraction. I tested the hypotheses that females would choose males based on their frequency and placement of scent marks, and that a female would advertise interest in a particular male by placing her scent marks on or near those of a preferred mating partner. In a series of experiments conducted with prairie voles, Microtus ochrogaster, females did not choose mates based on the frequency or placement of scent marks by males nor did they advertise their interest in a particular male through the frequency or placement of scent marks. The number of males chosen that scent-marked more than their opponents did not differ significantly between females exposed (11 of 15) and not exposed (10 of 15) to scents of males. Females exposed and not exposed to scents of males preferred seven of the same males that had scent-marked more than their opponents. When a third group of females was exposed to four times more scent of the less preferred than preferred males, they still chose the preferred males. Thus, the frequency and placement of scent marks by males were not used to assess males for mate choice nor did female prairie voles use scent to advertise their preference for a mating partner. In that scent marking is common in male and female mammals, scent quality might be more important than quantity in male-male competition and mate attraction.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Home range formation in wolves due to scent marking

Social carnivores, such as wolves and coyotes, have distinct and well-defined home ranges. During the formation of these home ranges scent marks provide important cues regarding the use of space by familiar and foreign packs. Previous models for territorial pattern formation have required a den site as the organizational center around which the territory is formed. However, well-defined wolf home ranges have been known to form in the absence of a den site, and even in the absence of surrounding packs. To date, the quantitative models have failed to describe a mechanism for such a process. In this paper we propose a mechanism. It involves interaction between scent marking and movement behavior in response to familiar scent marks. We show that the model yields distinct home ranges by this new means, and that the spatial profile of these home ranges is different from those arising from the territorial interactions with den sites.