The Social Behavior of Brown Spider Monkeys (Ateles hybridus) in a Fragmented Forest in Colombia

Forest fragmentation and habitat loss are two of the main drivers of wildlife population declines. Animals exposed to habitat disturbances must develop behavioral strategies to adapt to novel, rapidly fluctuating socioecological challenges. Understanding the behavioral flexibility of endangered primates as a response to ecological challenges, e.g., anthropogenic habitat disturbance, is a key element in the design of successful conservation initiatives for remaining populations. We studied the social behavior of a group of 11 adult and subadult brown spider monkeys (Ateles hybridus) living in a recently isolated and densely populated forest fragment in the Magdalena River Valley, Colombia, and compared their behavior to that of other wild spider monkey populations. From June 2009 to July 2010 we assessed diet, activity budgets, and the rates of affiliative and agonistic interactions initiated and received by adult male and female spider monkeys. The diet of our focal group was quite different from that in most previous studies of Ateles: leaves accounted for ca. 40?% of their diet, and fruits represented barely half of their diet, suggesting that this population has had to adjust its feeding strategies to the local ecological challenges. We found no sex differences in the time males and females spent resting, moving, and socializing, but females tended to invest more time in feeding than males did. Male-to-female aggression was the most common agonistic interaction, and same-sex aggressions were almost absent. We found no significant differences in rates of grooming or embracing between the different types of dyads. The resemblance of our results to those of similar studies in less disturbed habitats suggests that spider monkeys might adapt to certain levels of ecological and social disturbance, at least initially, and are a starting point to understand better the initial effects of fragmentation on the behavioral repertoire of these primates.     

Positional Behavior of Black Spider Monkeys (Ateles paniscus) in French Guiana

Spider monkeys (Ateles) frequently use suspensory locomotion and postures, and their postcranial morphology suggests convergence with extant hominoids in canopy and food utilization. Previous studies of positional behavior in Ateles, have produced variable rates in the use of different positional activities. I investigated the positional behavior of black spider monkeys (Ateles paniscus) in a wet rain forest in French Guiana, and assessed differences in the rates of use of positional modes across studies. I also discuss the significance of suspensory activities in forest utilization. In French Guiana, Ateles confined travel and feeding locomotion on small and medium-sized moderately inclined supports in the main canopy. Tail-arm brachiation and clamber were their main traveling modes, while clamber was the dominant feeding locomotor mode. Small horizontal supports were predominant during their feeding. Suspensory postures accounted for more than half of feeding bouts, with tail-hang and tail-hind limb(s) hang being the dominant postures. Feeding occurred largely in tree crown peripheries with the prehensile tail anchored frequently above the monkey. They usually collected food items below or at the same level as the body. There is no difference among the postures they used to acquire and eat young leaves and fruit. My results agree with reports on the positional behavior of different species of spider monkeys at other sites. Despite the use of different methods, the same species exhibited more or less similar profiles in similar forests. Interspecific differences could be associated with morphological differences. Moreover, intraspecific differences could be attributed to forest structure. The findings suggest that the major part of biological information is independent of methods used in the several studies. Suspensory behavior facilitates the exploitation of the forest canopy by shortening traveling pathways between and within trees, by enabling faster travel for the better exploitation of patchy food sources and by providing access to food in the flexible terminal twigs.     

Massive Destruction of Symphonia globulifera (Clusiaceae) Flowers by Central American Spider Monkeys (Ateles geoffroyi)1

Although more than 39 primate species have been described as nectar feeders, no studies have documented the negative affect this behavior may have on the reproductive success of the plants they consume. Here we report, for the first time, massive flower destruction of the tropical tree Symphonia globulifera by the frugivorous spider monkey, Ateles geoffroyi, and document the detrimental effect this behavior has on fruit set. Foraging behavior was collected from one troop of A. geoffroyi during 460 contact hours from June 1999 to May 2000 in the tropical humid forest at the Refugio de Vida Silvestre Punta Rio Claro (8°39′N, 83°44′E) in the Osa Peninsula in Southwestern Costa Rica. Detailed phenological data were collected from ten S. globulifera trees that the monkeys fed upon and ten trees that were outside of their home range, but in the Refugio. From July to September 1999 S. globulifera was the most important species consumed, representing from 86 to 100 percent of the total feeding time each month. Monkey foraging on S. globulifera flowers was destructive, detaching the flowers completely from the branches in 80 percent of the foraging bouts, and leaving flowers without petals and with damaged reproductive parts in 20 percent of the bouts. None of the ten trees where the monkeys were foraging set fruit, but seven of ten trees outside of their home range set fruit in November and December 1999. These results suggest that some primates may play an important role in floral herbivory in tropical forests and the subsequent reduction in reproductive success. Future studies should focus on evaluating the long‐term effect this foraging behavior may have on the population structure of this species and evaluate the effect of primate floral herbivory on the reproductive success of other plant species.     

Diet of the Critically Endangered Brown Spider Monkey (Ateles hybridus) in an Inter‐Andean Lowland Rainforest in Colombia

Brown spider monkeys (Ateles hybridus) are one of the least known and more threatened primates in the Neotropics. Recognized as a species about a decade ago, field studies on these endangered primates have mainly focused on estimating local population densities. Since 2006, we habituated a group of wild brown spider monkeys at Serranía de Las Quinchas, Colombia, and studied their feeding ecology during 2.5 years using focal “subgroup” sampling, and conducted phenological surveys in order to estimate habitat‐wide fruit availability. Based on 847 hr of behavioral follows, brown spider monkeys spent approximately 25% of their time in feeding activities, and fed from fruits and leaves on at least 123 plant species. Ripe fruits were the most important item in the diet of A. hybridus at Las Quinchas comprising 92% of their feeding time. Probably due to the minor variation in the monthly proportion of fruits in brown spider monkey's diet throughout this study, there was no relation between habitat‐wide fruit availability and the proportion of fruit included in their monthly diet. The diet of brown spider monkeys at Las Quinchas is toward the high end of fruit intake, even within other wild spider monkeys’ populations, suggesting that these endangered primates might also be facing the challenges of being a large bodied fruit specialist under a regional scenario of habitat loss and fragmentation. Am. J. Primatol. 74:1097‐1105, 2012. © 2012 Wiley Periodicals, Inc.     

Grooming,kinship, and co‐feeding in captive spider monkeys (Ateles geoffroyi)

Allo‐grooming is perhaps the most powerful affiliative behavior observed in non‐human primates. However, the functional significance of grooming in New World monkeys has not yet been fully understood, perhaps because grooming is less frequently observed in platyrrhines. To differentiate the potential role of affiliative investment and/or kinship on sharing access to food (co‐feeding) in spider monkeys, behavioral data on grooming, embracing, and feeding were collected from two different groups of captive study subjects: a familiar/kin group and a non‐familiar/non‐kin group. The results of this study suggest that family‐related spider monkeys that engage in grooming tend to share access to food resources more than unfamiliar conspecifics that do not groom. One explanation for this difference is that the unfamiliar study subjects had not yet invested in the affiliative social network, were not reciprocating their affiliative investments and hence, had a higher tendency toward single animal monopolizing resources. Degree of relatedness alone was not found to be a determinant for sharing the access to food, suggesting that familiarity in spider monkeys is based on the extent to which animals invest in affiliative relationships. In this study, only animals that had engaged in long‐term grooming and recognized each other as familiar shared the access to food. Therefore, it might be likely that in spider monkeys, long‐term grooming of high intensity has to be developed for co‐feeding to occur. Zoo Biol 20:293–303, 2001. © 2001 Wiley‐Liss, Inc.     

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.     

Effectiveness of Spider Monkeys (<Emphasis Type="Italic">Ateles geoffroyi vellerosus</Emphasis>) as Seed Dispersers in Continuous and Fragmented Rain Forests in Southern Mexico

Seed dispersal is considered a key process determining spatial structure and dynamics of plant populations, and has crucial implications for forest regeneration. We evaluated the effectiveness of spider monkeys (Ateles geoffroyi) as seed dispersers in continuous and fragmented habitats to test if this interaction is altered in forest fragments. We documented fruit and seed handling, defecation patterns, diversity and composition of seeds in feces, and seed germination of defecated and control seeds in the Lacandona rainforest, Mexico. For most species contributing to 80% of total fruit feeding time, monkeys swallowed and spat seeds, but swallowing was the most frequent seed handling category in continuous and fragmented forests. However, the proportion of feeding records of swallowed seeds was higher in continuous forest (0.59) than in fragments (0.46), whereas the opposite was true for proportion of dropped seeds (0.16 vs. 0.31). This pattern was reflected in the number of fecal samples containing seeds, which was greater in continuous (95.5%) than in fragmented forests (82.5%). Seeds in fecal samples included a total of 71 species from 23 plant families. The numbers of defecated seed species were similar between forest conditions, and in both cases most seeds (>86%) were undamaged. Defecated seeds showed greater germination percentages than control seeds in all of the five species evaluated. Although we identified some differences in seed handling and the percentage of feces with seeds between continuous forest and fragments, our results indicate that, in general terms, spider monkeys are effective seed dispersers in both forest conditions.     

Seasonal variation in association patterns of wild spider monkeys (<Emphasis Type="Italic">Ateles belzebuth belzebuth</Emphasis>) at La Macarena,Colombia

Spider monkeys exhibit a fission–fusion type of social organization. I studied party size and party composition in wild long-haired spider monkeys (Ateles belzebuth belzebuth) in three study periods at La Macarena, Colombia and found that overall party size was larger in the fruit-abundant season. Mean party size in which males were observed was relatively stable across seasons. In contrast, the mean party size of females varied. Females were observed in larger parties in the fruit-abundant season than in the fruit-scarce season. Moreover, whereas males associated with each other at an almost equal frequency across seasons, females associated with each other more frequently in the fruit-abundant season. Females with infants or small juveniles were more often in association with other individuals than were cycling females. The intensity of individual relationships varied according to season, such that even mothers and sons were not always strongly associated. In a large party, females with infants may gain from predation avoidance but they are at a disadvantage in terms of scramble competition. The balance between these factors may change with fruit availability and may influence party size in different periods. For males, party formation may facilitate the defense of resources from neighboring groups more than provide predation avoidance. Electronic Publication     

Housing with Females Increases Testosterone and Cortisol Levels in Captive Groups of Black-Handed Spider Monkeys (Ateles geoffroyi)

The black‐handed spider monkey (Ateles geoffroyi) is a seasonal reproducer that requires a seclusiveness to copulate and has a fusion–fission social system. These features impose important restrictions to achieve reproduction of captive animals. We investigated if group composition in captive spider monkeys has any endocrine effects. We compared testosterone and cortisol concentrations during the mating season in all‐male and multifemale–multimale groups to study if the former condition impairs reproductive potential and increases stress. Concentrations of testosterone and cortisol of males living with females were higher than those of all‐male groups. In the multifemale–multimale condition, dominant males had the highest levels of testosterone, while the youngest males showed the highest concentrations of cortisol. Results show that males adjust well to isosexual grouping, this being an appropriate condition to keep animals when controlled reproduction is sought. Zoo Biol 31:490;–497, 2012. © 2012 Wiley Periodicals, Inc.     

Seed Dispersal by Monkeys and the Fate of Dispersed Seeds in a Peruvian Rain Forest1

Primary seed dispersal by two species of monkeys and the effects of rodents and dung beetles on the fate of dispersed seeds are described for a rain forest in southeastern Perú. During the six-month study period (June–November 1992) spider monkeys (Ateles paniscus) dispersed the seeds of 71 plant species, whereas howler monkeys (Alouatta seniculus) dispersed seeds of 14 species. Spider and howler monkeys also differed greatly in their ranging behavior and defecation patterns, and as a consequence, produced different seed rain patterns. Monkey defecations were visited by 27 species of dung beetles (Scarabaeidae). Dung beetles buried 41 percent of the seeds in the dung, but the number of seeds buried varied greatly, according to seed size. Removal rates of unburied seeds by rodents varied between 63–97 percent after 30 d for 8 plant species. The presence of fecal material increased the percentage of seeds removed by seed predators, but this effect became insignificant with time. Although seed predators found some seeds buried in dung balls (mimicking burial by dung beetles), depth of burial significantly affected the fate of these seeds. Less than 35 percent of Brosimum lactescens seeds buried inside dung balls at a depth of 1 cm remained undiscovered by rodents, whereas at least 75 percent of the seeds escaped rodent detection at a depth of 3 cm and 96 percent escaped at 5 cm. Both dung beetles and rodents greatly affected the fate of seeds dispersed by monkeys. It is thus important to consider postdispersal factors affecting the fate of seeds when assessing the effectiveness of frugivores as seed dispersers.     

Determination of vertebral heart score in three species of Spider monkey (Ateles fusciceps,A. hybridus and A. paniscus)

Background

The vertebral heart score (VHS) is a method of evaluation of cardiac size well documented in domestic mammals and in other primate species, and the aim of this study was to determine the VHS in three species of Spider monkey.

Methods

In this retrospective study, right lateral radiographs of thirty clinically well animals were reviewed and VHS determined. The species included were Ateles fusciceps (n=17), Ateles hybridus (n=8) and Ateles paniscus (n=5).

Results

The VHS was found to vary between species and was 9.73±0.81 for A. fusciceps, 10.53±0.37 for A. hybridus and 10.45±0.27 for A. paniscus.

Conclusions

The observed values appear consistent with values determined for other primate species. There was statistically significant variation noted between species, and so VHS should be considered species‐specific in this genus. The values determined may be of benefit in objectively evaluating cardiac size in the species investigated.     

闭弯尾姬蜂与菜蛾盘绒茧蜂寄生菜蛾幼虫时的种间竞争

在室内25℃下,以菜蛾3龄初幼虫作寄主,研究了菜蛾盘绒茧蜂Cotesia plutellae和半闭弯尾姬蜂Diadegma semiclausum的种间竞争。当寄主供2种蜂同时产卵寄生时,2种蜂各自的寄生率与其单独寄生时无显著差异,合计寄生率比一种蜂单独存在时有所提高,但差异不显著。2种蜂均能产卵寄生已被另一种蜂寄生了的寄主幼虫。当寄主被2种蜂寄生的间隔时间很短（少于10 h）时,所育出的蜂绝大部分（80%以上）为绒茧蜂;当寄主先被绒茧蜂寄生,并饲养2天以上再供弯尾姬蜂寄生时,所育出的全为绒茧蜂;当寄主先被弯尾姬蜂寄生,并饲养2天以上再供绒茧蜂寄生时,寄主幼虫绝大部分不能存活,只有少部分能育出寄生蜂,且多为弯尾姬蜂。当2种蜂的幼虫存在于同一寄主体内时,2种蜂的发育均受到另一种蜂的抑制;绒茧蜂1龄幼虫具有物理攻击能力,能将弯尾姬蜂卵或幼虫致死。这些结果表明,菜蛾盘绒茧蜂与半闭弯尾姬蜂在同一寄主中发育时,前者具有明显的竞争优势。     

荒漠草原三种蝗虫成虫种内和种间竞争的研究

荒漠草原三种蝗虫成虫种内和种间竞争的研究贺达汉方成郑哲民（宁夏农学院农学系,永宁７５０１０５）（陕西师范大学动物研究所,西安７１００６２）Ｉｎｔｒａ＿ａｎｄＩｎｔｅｒｓｐｅｃｉｆｉｃＣｏｍｐｅｔｉｔｉｏｎａｍｏｎｇｔｈｅＡｄｕｌｔｓｏｆＴｈｒｅｅＧｒ...     

Population genetic patterns among social groups of the endangered Central American spider monkey (Ateles geoffroyi) in a human‐dominated landscape

Spider monkeys (Genus: Ateles) are a widespread Neotropical primate with a highly plastic socioecological strategy. However, the Central American species, Ateles geoffroyi, was recently re‐listed as endangered due to the accelerated loss of forest across the subcontinent. There is inconsistent evidence that spider monkey populations could persist when actively protected, but their long‐term viability in unprotected, human‐dominated landscapes is not known. We analyzed noninvasive genetic samples from 185 individuals in 14 putative social groups on the Rivas Isthmus in southwestern Nicaragua. We found evidence of weak but significant genetic structure in the mitochondrial control region and in eight nuclear microsatellite loci plus negative spatial autocorrelation in Fst and kinship. The overall pattern suggests strong localized mating and at least historical female‐biased dispersal, as is expected for this species. Heterozygosity was significantly lower than expected under random mating and lower than that found in other spider monkey populations, possibly reflecting a recent decline in genetic diversity and a threat from inbreeding. We conclude that despite a long history of human disturbance on this landscape, spider monkeys were until recently successful at maintaining gene flow. We consider the recent decline to be further indication of accelerated anthropogenic disturbance, but also of an opportunity to conserve native biodiversity. Spider monkeys are one of many wildlife species in Central America that is threatened by land cover change, and an apt example of how landscape‐scale conservation planning could be used to ensure long‐term persistence.     

Maintenance of female dominance in blue‐eyed black lemurs (Eulemur macaco flavifrons) and gray bamboo lemurs (Hapalemur griseus griseus) under semi‐free‐ranging and captive conditions

Several species of Malagasy prosimians are characterized by female dominance, an unusual trait among mammals. We compare the extent to which female dominance is displayed and the mechanisms that are used to maintain dominance in the frugivorous blue‐eyed black lemurs (Eulemur macaco flavifrons) and the folivorous gray bamboo lemurs (Hapalemur griseus griseus) housed at the Duke Lemur Center. All dominant–submissive interactions were recorded during 448 hr of focal animal observations. Both species of lemurs exhibited clear patterns of female dominance. However, the two species used aggressive dominance (defined as aggression+submission) and social dominance (defined as submission in the absence of aggression) to different extents in maintaining hierarchies within each group. The adult female blue‐eyed black lemurs used aggressive dominance (e.g., chase, cuff, bite) in more of their dominance interactions (66%) than did the adult female gray bamboo lemurs (40%). In both species, rates of aggressive dominance interactions were higher during feeding versus nonfeeding periods and while in smaller outdoor runs versus larger natural habitat enclosures, but the differences were not significant. Overall blue‐eyed black lemurs exhibit a more aggressive form of female dominance compared to the gray bamboo lemur. Zoo Biol 26:345–361, 2007. © 2007 Wiley‐Liss, Inc.     

Social Reversal of Sex‐Biased Aggression and Dominance in a Biparental Cichlid Fish (Julidochromis marlieri)

In biparental species, aggression, dominance, and parental care are typically sexually dimorphic. While behavioral dimorphism is often strongly linked to gonadal sex, the environment—either social or ecological—may also influence sex‐biased behavior. In the biparental cichlid fish Julidochromis marlieri, the typical social environment for breeding pairs consists of large females paired with smaller males. While both sexes are capable of providing territory defense and parental care, the larger female provides the majority of defense for the pair, while the smaller male remains in the nest guarding their offspring. We examine the contributions of sex and relative mate size to these sex‐biased behaviors in monogamous J. marlieri pairs. Both female‐larger and male‐larger pairs were formed in the laboratory and were observed for territorial aggression (against conspecifics and heterospecifics), dominance, and parental care. In female‐larger pairs, territorial aggression and intra‐pair dominance were female‐biased, while in male‐larger pairs this bias was reversed. For both pairing types, the presence of an intruder amplified sex differences in territorial aggression, with the larger fish always attacking with greater frequency than its mate. Though less robust, there was evidence for plasticity of sex‐bias for some egg care related behaviors in the inverse direction. Our study suggests that relative mate size strongly influences the sex bias of aggression and dominance in J. marlieri and that this aspect of the social environment can override the influence of gonadal sex on an individual's behavior. The remarkable plasticity of this species makes Julidochromis an exciting model that could be used to address the relationship between proximate and ultimate mechanisms of behavioral plasticity.     

The Association of Intergroup Encounters,Dominance Status,and Fecal Androgen and Glucocorticoid Profiles in Wild Male White‐Faced Capuchins (Cebus capucinus)

Androgens play a role in male reproductive competition, frequently via aggression, while glucocorticoids are associated with the stress response. However, the relationships of these hormones with different sources of competition (intra‐ vs. intergroup) and dominance status are highly variable. Here, we consider the fecal androgen (fA) and glucocorticoid (fGC) profiles of alpha and subordinate male Cebus capucinus in the context of intergroup competition during a rare period of low intragroup competition (i.e. all females were either pregnant or lactating). Intergroup encounters (IGEs) are a long‐term reproductive strategy in male white‐faced capuchins, enabling them to assess the composition of neighboring groups. IGEs pose a threat to resident males as these can result in injury or death, loss of dominance rank, group eviction, and group takeovers that are frequently associated with infanticide. From February to July 2007, fecal samples were collected from eight males in three groups of white‐faced capuchins in the Santa Rosa Sector of the Área de Conservación Guanacaste, Costa Rica. IGE rate was positively associated with both fA and fGC levels, indicating that IGEs are perceived as reproductive challenges by resident males, and may be associated with elevated metabolic costs. Alpha males sire the majority of group offspring and, accordingly, the threat of IGEs to both future (via rank loss or eviction) and current (via infanticide) reproductive success is greater than for subordinate males. Consistent with this observation, alpha males had higher fA and fGC levels than subordinate males. Given that all females were either pregnant or lactating and pronounced overt intragroup competition was absent, we interpret the difference in hormone profiles of alpha and subordinate males as being primarily associated with variation in the perceived threats of IGEs according to dominance status. Future studies should focus on the interaction of intra‐ and intergroup competition by examining hormone levels in the presence of periovulatory females. Am. J. Primatol. 75:107‐115, 2013. © 2012 Wiley Periodicals, Inc.     

Contest‐type competition between age classes in scramble‐type Callosobruchus maculatus (Coleoptera: Bruchidae)

We examined the effect of age differences on competition type in individuals of a scramble‐type strain of Callosobruchus maculatus (F.). When oviposition of two individuals on a bean was manipulated to introduce time intervals using two lines with different adult body colors, the frequency of two‐adult emergence decreased with the introduction of sequential oviposition. This result indicates that an age difference between two individuals induces contest competition. The frequency of adult emergence in older individuals decreased, whereas in younger individuals it increased with the introduction of sequential oviposition. Using a dissecting microscope, we observed that bodies of older individuals that died in the bean during the 4‐day oviposition interval were crushed at the pupal stage under the pupal chambers of younger individuals. These results show that an age difference between two larvae in a bean causes contest competition due to one‐sided interference by a younger individual during pupation of an older individual. Based on these experimental results, we discuss the ecological cause of contest competition and the population‐level consequences of identified interactions in scramble‐type C. maculatus.     

No short‐term change in avian assemblage following removal of Yellow‐throated Miner (Manorina flavigula) colonies

The overabundance of Yellow‐throated Miner (Manorina flavigula) has been shown to negatively affect the abundance and richness of small birds in areas they occupy, leading to homogenization of the avifauna across the fragmented landscape. In this study, we took advantage of a planned management cull to ask the question, does the removal of Yellow‐throated Miner colonies cause an immediate change in avian species richness and abundance? This cull was undertaken around the Bronzewing Flora and Fauna Reserve (north‐western Victoria, Australia) in order to protect a resident population of endangered Black‐eared Miner (M. melanotis) from hybridization. We conducted avian surveys along roadsides surrounding the reserve at Yellow‐throated Miner colonies (n = 6), control sites with no miners (n = 7), and where colonies were removed (n = 3). We found that the cull was followed by only a very modest increase in the species richness and abundance of small birds, with no significant effects on avian assemblage overall. This result contrasts with far more dramatic increases following culls of other species of miner. Sites where miners were removed were not depauperate of other species prior to the cull, which could have been due to a combination of proximity to refuge for small birds in a neighbouring reserve or the low numbers of miners that made up each culled colony. This study highlights that assumed effects of a management action may be highly dependent upon spatial and temporal context.