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Many spring-feeding insect folivores have evolved to hatch in synchrony with temporal windows of high foliar quality that occur during leaf expansion. In some host-tree species, however, budbreak phenology can shift in response to defoliation during the prior growing season. While this response could have important impacts on the performance of insect folivore species prone to multi-year outbreaks, the prevalence, underlying mechanisms, and ecological ramifications of defoliation-induced phenological shifts are poorly understood. We established a common garden of twelve trembling aspen (Populus tremuloides Michx.) genotypes, and observed the effects of prior-season defoliation and soil-nutrient availability on budbreak phenology during the following spring. Average budbreak phenology was delayed in response to both prior-season defoliation and low soil-nutrient treatments, but these responses were lower in magnitude than those previously shown to occur in response to temperature. Magnitudes of budbreak delay varied significantly among genotypes and were highest among early-flushing aspen. This negative correlation between aspen budbreak phenology and magnitudes of defoliation-induced phenological delay suggests that widespread defoliation could decrease intraspecific variation in aspen budbreak phenology during subsequent growing seasons, potentially allowing larvae to adapt to windows of optimal foliage quality during multi-year outbreaks. These results quantify additional environmental factors that may act in concert with major drivers such as temperature to mediate phenological relationships between aspen and folivorous insects.

Forest cover is decreasing globally, chiefly due to the conversion of forest to agricultural landscapes. In contrast, the area under plantation forestry is increasing by up to 3 million hectares per annum. For wildlife occupying landscapes where native forest is the dominant land cover, plantations generally represent a lower value habitat; however, plantations established on land formerly used for pasture, may benefit wildlife by providing temporary forest habitat and increasing connectivity. This study investigates the influence of landscape, site and climatic factors on koala population density in far south-west Victoria where there has been extensive plantation establishment. We conducted koala surveys and habitat characteristic assessments at 72 sites across three habitat types: plantation, native vegetation blocks, and native vegetation strips. We employed a hierarchical modelling framework for estimating abundance, and constructed candidate multinomial N-mixture models to identify factors influencing the abundance of koalas. We detected higher mean koala density in plantation sites (0.85 per ha) than in either native block (0.68 per ha) or native strip sites (0.66 per ha). We found five covariates of koala density, and using these variables we spatially modelled koala abundance and discuss factors that are key in determining large-scale distribution and density of koala populations. We provide a distribution map that can be used to identify high priority areas for population management as well as habitat of high conservation significance for koalas. This information facilitates the linkage of ecological theory with the on-ground implementation of management actions, and may guide conservation planning and resource management actions to consider overall landscape configuration as well as the spatial arrangement of plantations adjacent to remnant forest.

For species that are increasingly threatened by the combined impacts of habitat loss and climate change, the identification of priority regions for conservation planning efforts is urgently required. In the case of specialist folivores, consideration of the effects of climate change on the distributions of their essential food resources should be a key component of conservation planning. The koala (Phascolarctos cinereus) was listed in 2012 as vulnerable under Australian Commonwealth Government law. Here, we incorporate species distribution models for the koala, an arboreal marsupial, and its specialized food resources, to identify broad-scale priority conservation regions. We demonstrate a spatial prioritization approach that informs conservation planning and that links the shifting distribution of this declining species and its critical food resources under climate change. We find that the inclusion of food plants affected the identification of priority regions for conserving koalas, and that priority regions for the conservation of this species are predicted to shift considerably, often outside the current range of this species, posing additional challenges for its conservation.

The gypsy moth (Lymantria dispar) is an insect folivore that feeds on a broad range of hosts, and undergoes intermittent outbreaks that cause extensive tree mortality. Like many other herbivorous insects, gypsy moth larvae consume a substrate that is low in nitrogen. Gypsy moth larvae have been known to cannibalize under crowded conditions in the laboratory. In this study, we assessed the influence of nitrogen and density on cannibalism behavior in gypsy moth larvae. Cannibalism rates increased with decreased nitrogen and increased density. There was no interaction between these two parameters. Developmental experiments confirmed that low dietary nitrogen is detrimental, in agreement with previous studies. In a second experiment, we assessed the influence of previous cannibalism experiences on subsequent cannibalism behavior. Gypsy moth larvae that had previously cannibalized other larvae subsequently exhibited higher cannibalism rates than those larvae that had not cannibalized. In conclusion, low nitrogen, high larval density, and previous cannibalism experience are important factors contributing to gypsy moth larval cannibalism. Future studies are needed to estimate benefits to larvae, and to more closely approximate field conditions.

The infraorbital foramen (IOF) is located below the orbit and transmits the sensory infraorbital nerve (ION) to mechanoreceptors located throughout the maxillary region. The size of the IOF correlates with the size of the ION; thus, the IOF appears to indicate relative touch sensitivity of maxillary region. In primates, IOF size correlates well with diet. Frugivores have relatively larger IOFs than folivores or insectivores because fruit handling/processing requires increased touch sensitivity. However, it is unknown if the IOF can be used to detect subtle dietary differences among closely related hominoid species. Hominoids are traditionally grouped into broad dietary categories, despite the fact that hominoid diets are remarkably diverse. This study examines whether relative IOF size is capable of differentiating among the dietary preferences of closely related species with overlapping, yet divergent diets. We measured IOF area in Hylobates lar, Symphalangus syndactulus, Pongo pygmaeus spp., Pan troglodytes, Gorilla gorilla, Gorilla beringei graueri, and Gorilla beringei beringei. We classified each species as a dedicated folivore, mixed folivore/frugivore, soft object frugivore, or hard object frugivore. The IOF is documented to be larger in more frugivorous species and smaller in more folivorous taxa. Interestingly, G.b. beringei, had the largest relative IOF of any gorilla, despite being a dedicated folivore. G.b. beringei does have unique food processing behavior that relies heavily on maxillary mechanoreception, thus this finding is not entirely unsupported behaviorally. The results of this study provide evidence that the IOF is an informative feature in interpretations of fossil apes. Anat Rec, 297:1377–1384, 2014. © 2014 Wiley Periodicals, Inc.

Landscape disturbance can cause shifts in vegetation communities, which can impact animal populations. However, the processes responsible for the cascading effects between plants and animals are often poorly understood. We demonstrate a mechanism by which landscape disturbance can impact habitat quality by altering the nutritional composition of a forest using a case-study of koalas (Phascolarctos cinereus) in Australian eucalypt forests. We used two large datasets, a detailed survey of forest tree species composition and koala occurrence across 26,000 ha of forests on the South East Coast of Australia and an extensive eucalypt nutritional quality database, to investigate relationships among tree species composition, nutritional quality, and koala occurrence. Eucalyptus sieberi, a species native to the study area, responds well to disturbance and can increasingly dominate forests after logging and/or fire. Using negative binomial models, we found that observations of koala scat decreased as the proportion of E. sieberi in an area increased beyond zero. Eucalyptus sieberi had lower available nitrogen (AvailN) and total nitrogen than most other eucalypt species, and the highest concentrations of unsubstituted B-ring flavanones, an herbivore-deterrent plant secondary metabolite for the koala. Simulations of forests with increasing proportions of E. sieberi showed that AvailN decreased from 1.1% when E. sieberi was ≤10% of all eucalypts to 0.7% AvailN when E. sieberi was the only species present. Consequently, forests dominated by E. sieberi are unlikely to support koala populations based on existing knowledge of koala nutritional requirements. Foliage nutritional quality is a critical but often overlooked element of habitat quality for leaf eating animals, and this study demonstrates how disturbance related changes in tree species composition can impact the nutritional landscape and the occurrence of folivores.

Despite the essential role of Bifidobacterium in health-promoting gut bacteria in humans, little is known about their functions in wild animals, especially non-human primates. It is difficult to determine in vivo the function of Bifidobacterium in wild animals due to the limited accessibility of studying target animals in natural conditions. However, the genomic characteristics of Bifidobacterium obtained from the feces of wild animals can provide insight into their functionality in the gut. Here, we analyzed the whole genomes of 12 B. moukalabense strains isolated from seven feces samples of wild western lowland gorillas (Gorilla gorilla gorilla), three samples of wild central chimpanzees (Pan troglodytes troglodytes) and two samples of wild forest elephants (Loxodonta cyclotis) in Moukalaba-Doudou National Park, Gabon. In addition, we analyzed the fecal bacterial communities of six wild western lowland gorillas by meta 16S rRNA gene analyses with next generation sequencing. Although the abundance of the genus Bifidobacterium was as low as 0.2% in the total reads, a whole genome analysis of B. moukalabense suggested its contribution digestion of food and nutrition of frugivore/folivore animals. Specifically, the whole genome analysis indicated the involvement of B. moukalabense in hemicellulose degradation for short chain fatty acid production and nucleic acid utilization as nitrogen resources. In comparison with human-associated Bifidobacterium spp., genes for carbohydrate transport and metabolism are not conserved in these wild species. In particular the glycosidases, which are found in all 12 strains of B. moukalabense, were variably detected, or not detected, in human-associated species.

The decisions that animals must make to achieve a balance between quantity and quality of resources become more difficult when their habitats are patchy and differ greatly in quality across space and time. Koalas are a prime subject to study this problem because they have a specialised diet of eucalypt leaves and need to balance nutrient and water intake against toxins in the leaves, all of which can change with soil type and climate. Koalas are nocturnal and spend most of the day resting and therefore choose trees for reasons other than feeding, particularly for thermoregulation. We GPS-tracked 40 koalas over 3 yr to determine their shift in tree selection between day and night, and in relation to daily maximum temperature, in a patchy rural landscape in north-western NSW, Australia. The species, degree of shelter, diameter, height and elevation of each visited tree were recorded. We used generalised linear mixed effects models to compare tree use between day and night and maximum daily temperature. Koalas used more feed-trees during the night, and more shelter-trees during the day. They also selected taller trees with more shelter in the day compared with night. As daytime temperatures rose, koalas increasingly selected taller trees at lower elevations. Our results demonstrate that koalas need taller trees, and non-feed species with shadier/denser foliage, to provide shelter from heat. This highlights the need both for the retention of taller, mature trees, such as remnant paddock trees, and the planting of both food and shelter trees to increase habitat area and connectivity across the landscape for arboreal species. Retaining and planting trees that provide optimum habitat will help arboreal folivores cope with the more frequent droughts and heatwaves expected with climate change.

In birch, Betula pubescens, herbivore-induced delayed induced resistance (DIR) of defoliated trees may cause a strong reduction in the potential fecundity of a geometrid folivore Epirrita autumnata. In this study, we examined the biochemical basis of DIR in birch leaves during a natural outbreak of E. autumnata. A set of experimental trees was defoliated at four sites by wild larvae in the peak year of the outbreak, whereas control trees were protected from defoliation by spraying with an insecticide. The biochemical composition of leaves was analysed in the following year and, although the DIR response was weak during this outbreak, causing less than a 20% reduction in the potential fecundity of E. autumnata, some consistent relationships between defoliation, biochemistry and pupal mass of E. autumnata suggested a general biochemical basis for the defoliation-induced responses in birch leaves. Total concentrations of nitrogen, sugars and acetone-insoluble residue (e.g. cell wall polysaccharides, cell-wall-bound phenolics, protein, starch, lignin and hemicellulose) were consistently lower, and total concentrations of phenolics, especially of gallotannins and soluble proanthocyanidins, were higher in the leaves of trees defoliated in the previous year than in those protected from defoliation. The capacity of tannins to precipitate proteins correlated with contents of gallotannins, and was highest in defoliated trees. The pupal mass of E. autumnata showed a strong, positive correlation with concentrations of nitrogen and sugars, and a negative correlation with the acetone-insoluble residue and gallotannins in foliage. Correlations with other measured biochemical traits were weak. The correlation coefficients between biochemical traits and pupal mass consistently had similar signs for both defoliated and insecticide-sprayed trees, suggesting that variation in leaf quality due to defoliation in the previous year was based on similar biochemical traits as variation for other reasons. We suggest that DIR is associated with reduced growth activity of leaves, and may be seen as a delay in the biochemical maturation of leaves in defoliated trees. This explains the high concentration of gallotannins in defoliated trees, a characteristic feature of young leaves. However, the lower content of nitrogen and the higher content of soluble proanthocyanidins in defoliated trees are traits usually characterising mature, not young, leaves, indicating defoliation-induced changes in chemistry in addition to modified leaf age. Our results emphasise the importance of understanding the natural changes in chemistry during leaf maturation when interpreting defoliation-induced changes in leaf biochemistry.

We examined the effects of the rhizome galling fly, Eurosta comma (Wiedemann), on rhizome mass and nitrogen allocation in a clone of its goldenrod host plant, Solidago missouriensis Nutt. In comparison to ungalled ramets, galled ramets initiated significantly fewer new rhizomes, and allocated less mass to leaves and stems and more to roots. Galled ramets had lower concentrations of nitrogen in roots and rhizomes but leaf and stem nitrogen concentrations were not affected. In the second year of our study, outbreaks of the leaf-eating beetle, Trirhabda canadensis (Kirby), occurred in three of our four study clones, removing ∼20%, ∼50%, and 100% of leaf area from clones 2–4, respectively. In the most heavily grazed clones, the influence of rhizome galls on plant resource allocation was least pronounced. Despite the overwhelming immediate impact of grazing, the presence of a gall population may be important because they decrease the ability of S. missouriensis clones to initiate new rhizomes and hence to recover from defoliation.

Estimating the nutritional value of a herbivore's diet is difficult because it requires knowing what the animal eats, the relative quality of each component and how these components interact in relation to animal physiology. Current methods are cumbersome and rely on many assumptions that are hard to evaluate. We describe a new method for estimating relative diet quality directly from faeces that avoids the problems inherent in other methods. We combine this method with near infrared reflectance spectroscopy (NIRS) to analyse many samples and thus provide a technique with immense value in ecological studies. The method stems from the correlation between the concentrations of dietary and faecal nitrogen in herbivores eating a tannin-free diet, but a weaker relationship in browsers that ingest substantial amounts of tannins, which form complexes with proteins. These complexes reduce the availability of nitrogen and may increase faecal nitrogen concentrations. Using the tannin-binding compound, polyethylene glycol, we showed that tannin-bound nitrogen is a significant and variable part of faecal nitrogen in wild common brushtail possums (Trichosurus vulpecula). We developed a technique to measure faecal available nitrogen and found that it predicted the reproductive success of female brushtail possums in northern Australia. Faecal available nitrogen combined with NIRS provides a powerful tool for estimating the relative nutritional value of the diets of browsing herbivores in many ecological systems. It is a better indicator of diet quality than other commonly used single-nutrient measures such as faecal nitrogen and foliage analysis paired with observed feeding behaviour.

Habitat loss and natural catastrophes reduce the resources available to animals. Species can persist if they have access to additional resources and habitats through the processes of landscape complementation and supplementation. In arid and semi-arid ecosystems, where productivity is limited by precipitation, the impact of landscape change and prolonged drought is severe on specialist species whose range boundaries are limited by aridity. We examined the pattern of occurrence by a specialist arboreal folivore, the koala, at the periphery of its biogeographic range, in a semi-arid rangeland landscape. We used hierarchical mixed modelling to examine the effect of landscape change on koala populations and their habitat use during and after a prolonged drought. We found that the tree species and the distance of a site from water courses were the most important determinants for koala presence in these landscapes. Koalas were predominantly detected in riverine habitat along the water courses, which are primary habitat and provide refugia in times of drought and extreme heat. There was a strong positive effect from the interaction between the amount of primary and secondary habitat in the landscape, although individually, the amount of each of these habitats was not important. This shows koalas will persist in more intact landscapes. There was no difference in habitat use between dry and wet years, but we consider that it can take several wet seasons for koalas to expand into habitats away from water courses.

The urbanization of the natural landscape has spread along the major axis of all large towns worldwide, what has resulted in an unprecedented increase in habitat fragmentation and ultimately in a mosaic of patches of native vegetation. These changes have a continuum of impacts on the processes and mechanisms that shape habitat diversity and species interactions. An important and overlooked aspect that may aid in the understanding of the effects of urbanization on species interactions is herbivory. Herbivores and leaf removal can signalize the effects of the habitat via their host plants. Another potentially useful measure for evaluating the effects of environmental stress on particular species is fluctuating asymmetry. We studied the herbivory levels and leaf fluctuating asymmetry on Solanum lycocarpum (Solanaceae) along an urban gradient in Brazil. The leaf area removed by folivores differed among sites and was significantly greater in the site with the higher level of urbanization. Also, the area removed varied significantly among individuals within urbanization sites. Similarly, the frequency of leaves with folivore damage was greater in the site with higher urbanization, although the area removed by them did not differ between shorter and larger plants. FA was significantly greater in the more urbanized sites in comparison with the other sites in the urbanization gradient and varied among individuals within site. Finally, we found a positive relationship between leaf area removed and FA values in the urbanization gradient. These results indicate that increased urbanization causes a reduction in habitat suitability for S. lycocarpum, which is reflected on higher FA levels. The leaf herbivores may benefit from the stress levels of hosts on more urbanized sites, which may have been caused by several factors acting in combination such as lower defenses, higher susceptibility and natural enemy pressures via habitat fragmentation/urbanization. This study exemplifies the variety of effects that urbanization exerts on remnant native populations and that fluctuating asymmetry could be a useful measure of the effects of these challenges on plant populations, and might also be used to assist management and conservation practices of natural areas within cities.

Edge effects are widely researched phenomena in ecology. However, little is known about their influence on the competition dynamics of sympatric species. We present a new conceptual model to explain how edge effects and competition dynamics interact to influence community composition and illustrate it with a case study of four sympatric arboreal marsupial species in Australia. For the first time, we demonstrate that edge effects on interspecific competition influence community composition in naturally co-occurring species. We found that Petauroides volans , a eucalypt folivore specialist, was significantly more edge-avoidant than the other three more generalist folivore species, Trichosurus vulpecula , Trichosurus cunninghami, and Pseudocheirus peregrinus . As the amount of edge increased, the abundance and probability of occurrence of P. volans decreased. We found the opposite pattern for the other species. We propose that edge effects differentially affect resource availability for these species and increase competitive pressure for tree hollows, which are a limited common resource.

Predicting variation in animal abundance across time and space has proven very difficult; however, a model exists to predict the biomass of small folivorous primates that has considerable correlative support. This model suggests that the protein-to-fiber ratio of leaves in a habitat can predict folivore biomass. Here we present an experimental test of this protein-to-fiber model to assess if the number of infant monkeys per female and group size can be predicted based on the leaf chemistry of a habitat. We expected regenerating forest in Kibale National Park, Uganda to have leaves with higher concentrations of crude protein and lower concentrations of fiber than old-growth forest trees, and consequently, we expected a greater number of infants per female in the folivorous red colobus ( Procolobus rufomitratus) with access to this area. As predicted, regenerating forests did have trees with leaves with high concentrations of protein and low concentrations of fiber, but there was no corresponding change in the demographic structure of red colobus groups. We also tested whether energy was a potential determinant of these parameters, but found no evidence for its importance. Our findings support recent studies that are critical of the protein-to-fiber model, which lead us to question the model's generality, particularly for conservation and management.

Body size is often associated with a dietary divergence within taxonomically related groups so that large animals are often folivorous, while smaller species shift progressively towards omnivory or carnivory. This trend may be influenced by allometric constrains which result in relatively high energetic requirements, but low gut capacities in small animals, compared to their large counterparts. The common brushtail possum (Trichosurus vulpecula, Phalangeridae) has six subspecies ranging widely in weight (1-4kg). They are not strictly folivorous, but supplement their diet with more nutritious, non-foliar foods. We predicted that T.vulpecula subspecies diverged in diet in association with body size, with smaller subspecies consuming higher proportions of non-foliar foods. We assessed this with a review and a meta-analysis of previous Australian studies. We also investigated the previously unquantified diet of T.v.hypoleucus at three sites in the northern jarrah forest of Western Australia. Results from the meta-analysis and the review supported our prediction. However, the large variability in the data highlighted their limitations and those of the techniques commonly used to quantify the diet of T.vulpecula. Nonetheless, small subspecies of T.vulpecula appear to consume higher proportions of non-foliar foods. These results should encourage further research into the body size/diet relationship within T.vulpecula and other possum species. Results from the dietary study of T.v.hypoleucus emphasized their omnivorous diet, which was dominated by foliage and flowers and smaller proportions of invertebrates, seeds and fruits. The common brushtail possum is seldom an exclusive arboreal folivore, but rather ranges from folivory to omnivory.

The correlation between the biomass of forest primates and a chemical index of the average nutritional quality of leaves in tropical forests has been repeatedly documented since 1990. We tested the role played by protein : fibre on lemur biomass in a gallery forest in southern Madagascar. Plant species abundance was determined based on transect censuses. We calculated an average ratio of protein-to-fibre in leaves and an abundance-weighted ratio, i.e. the mean weighted by the basal area of tree species, to be compared with the figures available for other forest ecosystems in Madagascar and a number of anthropoid habitats. Lemur densities were evaluated through compilation of previous studies made from prior to 1975 and up until 2011 based on strip censuses and/or identification of all groups supplemented with new censuses. A high mean ratio of protein to fibre (> 0.4) supports high folivore biomass at 390 kg km−2 (reaching 630 kg km−2 in the closed-canopy forest area) compared with primate communities in other Malagasy forests (protein : fibre: < 0.5; folivore biomass: < 440 kg km−2), as predicted. However, the data corroborate the finding that the total biomass of lemur communities as well as the biomass of folivorous lemur species are low compared with those of African and Asian primate communities for a given protein : fibre ratio. Tree diversity and leaf production do not consistently explain this pattern. In contrast, the extinction of large folivorous lemurs during the past two millennia presumably allowed too little time for smaller-sized species to evolve equally effective morphological and physiological specializations for processing a large range of fibrous foods.

Vigilance often decreases with increasing group size, due to lower predation risk or greater scramble competition for food. A group size effect on vigilance is seldom seen in primates, perhaps because scanning and feeding often occur simultaneously or because the distinction between routine and induced vigilance has not been investigated. We analyzed feeding and resting observations separately while distinguishing between routine and induced scans in four groups of wild ursine colobus monkeys (Colobus vellerosus) experiencing scramble competition for food and infanticide risk. We used linear mixed-effect models to test the effect of group size, age–sex class, number of neighbors, number of adult male neighbors, and height in the canopy on scanning rates (vigilance) with and without evident conspecific threat. Food type was also examined in the feeding models. Perceived predation risk affected vigilance more than scramble competition for food and infanticide risk. Routine and induced vigilance were greatest at lower canopy heights during feeding and resting and increased when individuals had fewer neighbors while resting. A group size effect was found on induced vigilance while resting, but scanning increased with group size, which probably indicates visual monitoring of conspecifics. Scanning rates decreased while feeding on foods that required extensive manipulation. This supports the idea that vigilance is relatively cost free for upright feeders when eating food that requires little manipulation, a common feature of folivore diets. In the presence of threatening conspecific males, close proximity to resident males decreased individual vigilance, demonstrating the defensive role of these males in the group.

A three-dimensional geometric morphometric approach was employed to examine shape variation in laser-scan generated models of lower second molars and its relationship to diet in a sample of 9 extant and 16 extinct platyrrhine genera. Principal component analysis of twenty-three x,y,z landmarks describing the occlusal table and sidewalls showed that dental relief was the main contributing factor to variation along the first axis. Discriminant function analysis (DFA) of PC 1 scores and centroid size accurately classified extant platyrrhines according to dietary preference; however, without centroid size, the DFA was less successful. Within this framework, most of the fossil platyrrhines, including specimens from Patagonia, Colombia, Brazil, and the Caribbean, were predicted to have had a frugivorous diet, but several taxa were classified as having a frugivorous/insectivorous diet, the middle Miocene Neosaimiri, Patasola, and Laventiana, all from La Venta. Alouattins, including the La Ventan Stirtonia and the Cuban Paralouatta, showed variable classification as either frugivores or folivore/frugivores. Xenothrix, from Jamaica, was classified either as a frugivore or frugivore/omnivore. Dietary profiles across different extinct platyrrhine communities are compared and discussed in a paleoecological context. Anat Rec, 294:2073–2091, 2011. V C 2011 Wiley Periodicals, Inc.

Activity and behavior patterns are important components of a given species’ ecological strategy, as they have profound implications for its survival and reproduction. Here, we studied the activities, movements and secretive behavior of the thin-spined porcupine Chaetomys subspinosus (Rodentia: Erethizontidae), a threatened arboreal folivore in the Brazilian Atlantic rainforest. We aimed to ascertain the behavioral strategies used by this species as well as its responses to seasonal and daily climatic changes. Four radio-collared individuals were followed continuously for 72-h in the summer and winter, as well as during 146 half-night sessions conducted from April 2005 to September 2006 in forest remnants in southern Bahia. The thin-spined porcupines were nocturnally active (17:30–05:40 h), with peaks in activity and movement from 19:00 to 20:00 h and 03:00 to 04:00 h. Animals followed a circadian rhythm of activity during both the summer and winter. During the diel cycle, porcupines spent 74% of their time resting, 14% feeding, 11% traveling and 2% performing other activities. Distance traveled during the diel cycle averaged 277.5 ± 117.9 m sd. The mean movement rate during the night was 21.6 ± 30.1 m/h sd. No significant changes in activity budget or daily distance traveled were observed between seasons, most likely in response to the low fluctuations in climatic conditions and food availability throughout the year in the study region. However, rainfall reduced the time that the animals spent on feeding activities and explained day-to-day differences in activity budgets. We also provide details about intraspecific interactions and defecation behavior. Our observations confirmed that thin-spined porcupines, similar to other folivorous species, present low activity levels and short daily movements, and have adopted various cryptic habits, such as nocturnality, a solitary lifestyle, the tendency to leave offspring alone most of the time and defecation in concealed latrines.

Little research has addressed the impacts of invasive-species establishment on native forest insect communities. Such information is lacking even for gypsy moth, Lymantria dispar (L.) (Lepidoptera: Erebidae), the most thoroughly studied invasive forest insect. We investigated the ecological impacts of gypsy moth on native species at sites in north-central Ontario, Canada, with and without significant histories of gypsy moth defoliation over a 2-year period. Patterns in native forest caterpillar communities are described using measures of species diversity and multivariate analysis. We documented a transition from low-level to dominant gypsy moth populations. Sites with different gypsy moth outbreak histories exhibited differences in rank-abundance distributions and dominance structures in the first year of the study; by the second year, gypsy moth was dominant at sites of both types irrespective of their previous defoliation history. Contrary to our predictions, we found that gypsy moth outbreak history had no significant effects on native caterpillar community diversity or structure. However, sites with currently high gypsy moth abundance demonstrated significant shifts in late-season caterpillar community structure. Our results suggest that observed community differences were due to the presence of a highly abundant folivore, and not to permanent shifts in the native community because of the introduction of an invasive species.

Numerous studies have been made of rodent population and community dynamics, especially in arid ecosystems. Most have centered on understanding how total and species-specific densities of rodents change over time. The standing biomass each species contributes also is important to the energy available to the mesocarnivore community. Although density and standing biomass are related, how they might differ and if those differences are of importance to community structure and function have not received much attention. We analyzed 12 years of rodent density and body mass data from the Chihuahuan Desert in northern Mexico. Data were collected yearly in spring and fall from radial livetrapping webs. Total density and biomass changed significantly in a parallel manner from year to year, and both were related to precipitation and percent cover of grass and forbs. Based on density and biomass, the rodent community was dominated by 2 or 3 principal species. However, on a species-specific level, the numerically dominant species was a small-bodied granivore (Chaetodipus nelsoni), and a large-bodied folivore (Neotoma albigula) dominated in biomass. As total density increased, the proportion contributed by dominant species decreased. As total biomass increased, the proportion in the 2 dominant species increased and accounted for approximately 80% of total biomass. Over the 12 years of the study, species distributions based on density showed no directional change. In contrast, biomass of the rodent community gradually concentrated in a single, large-bodied folivore, N. albigula. Although total density and biomass responded similarly to precipitation and plant productivity, considerable differences between these 2 characteristics existed in their species-specific contributions to and changes within the community. The significance of these differences relative to foraging strategies and variable feeding opportunities within the community is discussed.

The alarming rate of deforestation in Madagascar is driving some endemic primates to extinction. Surprisingly, anthropogenic habitat disturbance is not always deleterious. The effect of disturbance on lemur abundance may be related to diet, with frugivorous species more prone to population declines than folivores or insectivores. To test the effects of disturbance on lemur abundance and group size, we surveyed 2 sites within contiguous forest at Ranomafana National Park, 1 lightly disturbed primary forest (Vato) and 1 heavily logged forest (Tala). We quantified forest structure variables along 6 survey routes and conducted 68 diurnal and 42 nocturnal lemur surveys. Canopy closure, canopy height, and understory visibility were greater in Vato than in Tala. We encountered 2 frugivorous lemurs (Eulemur rufifrons, Varecia variegata) and 1 folivore (Avahi peyrierasi) significantly more frequently in Vato than in Tala, whereas the opposite was true for the insectivorous Microcebus rufus. Rates did not differ statistically for 1 frugivore (Eulemur rubriventer) and 2 folivores (Propithecus edwardsi, Hapalemur griseus). Comparisons across the 6 survey routes suggest that the abundance was heterogeneous within as well as between sites. Neither group size nor composition differed between sites. Encounter rates for Varecia variegata were positively related to canopy closure, and encounter rates for Avahi peyrierasi were positively related to canopy height. Encounter rates for Microcebus rufus were negatively related to canopy closure, height, and understory visibility. Similar to other studies, the results suggest that some lemurs, including folivores, may cope with anthropogenic disturbance better than others, including some frugivores. Lemur abundance is heterogeneous, though, even on small spatial scales.

The process of selecting certain desirable traits for plant breeding may compromise other potentially important traits, such as defences against pests; however, specific phenotypic changes occurring over the course of domestication are unknown for most domesticated plants. Cranberry (Vaccinium macrocarpon) offers a unique opportunity to study such changes: its domestication occurred recently, and we have access to the wild ancestors and intermediate varieties used in past crosses. In order to investigate whether breeding for increased yield and fruit quality traits may indirectly affect anti-herbivore defences, the chemical defences have been examined of five related cranberry varieties that span the history of domestication against a common folivore, the gypsy moth (Lymantria dispar). Direct defences were assessed by measuring the performance of gypsy moth caterpillars and levels of phenolic compounds in leaves, and indirect defences by assaying induced leaf volatile emissions. Our results suggest that breeding in cranberry has compromised plant defences: caterpillars performed best on the derived NJS98-23 (the highest-yielding variety) and its parent Ben Lear. Moreover, NJS98-23 showed reduced induction of volatile sesquiterpenes, and had lower concentrations of the defence-related hormone cis-jasmonic acid (JA) than ancestral varieties. However, induced direct defences were not obviously affected by breeding, as exogenous JA applications reduced caterpillar growth and increased the amounts of phenolics independent of variety. Our results suggest that compromised chemical defences in high-yielding cranberry varieties may lead to greater herbivore damage which, in turn, may require more intensive pesticide control measures. This finding should inform the direction of future breeding programmes.

Uncertainty in recovery times of tropical forests can lead to mismanagement, such as in setting inappropriate harvesting rates or failing to achieving conservation targets. We use long-term plot data (17 y) to estimate recovery times of separate forest compartments, which experienced different levels of timber extraction within Kibale National Park, Uganda. We estimate that structural recovery (basal area) of heavily logged and moderately logged compartments will take respectively 112 and 95 y, when compared to adjacent mature forest. Our data suggests that recovery in terms of species composition will take significantly longer. Our estimates of structural recovery are derived from rates of change of diameter at breast height and basal area measurements which have been used traditionally as indicators of forest growth and productivity. Our results suggest that the severity of the logging has an impact on the rate of recovery, with current recovery rates estimated at 0.32 m 2 ha −1 y −1 in a moderately logged compartment and 0.25 m 2 ha −1 y −1 in heavily logged areas, highlighting the possible benefits of reduced impact harvesting in increasing long-term yields. We investigate how some representatives of the wildlife community were affected by differential recovery times and find that recovery times of frugivorous primate's forest habitats were 2.5 times slower when compared with folivorous primates.

Conservation planning and population assessment for widely-distributed, but vulnerable, arboreal folivore species demands cost-effective mapping of habitat suitability over large areas. This study tested whether multispectral data from WorldView-3 could be used to estimate and map foliar digestible nitrogen (DigN), a nutritional measure superior to total nitrogen for tannin-rich foliage for the koala (Phascolarctos cinereus). We acquired two WorldView-3 images (November 2015) and collected leaf samples from Eucalyptus woodlands in semi-arid eastern Australia. Linear regression indicated the normalized difference index using bands “Coastal„ and “NIR1„ best estimated DigN concentration (% dry matter, R2 = 0.70, RMSE = 0.19%). Foliar DigN concentration was mapped for multi-species Eucalyptus open woodlands across two landscapes using this index. This mapping method was tested on a WorldView-2 image (October 2012) with associated koala tracking data (August 2010 to November 2011) from a different landscape of the study region. Quantile regression showed significant positive relationship between estimated DigN and occurrence of koalas at 0.999 quantile (R2 = 0.63). This study reports the first attempt to use a multispectral satellite-derived spectral index for mapping foliar DigN at a landscape-scale (100s km2). The mapping method can potentially be incorporated in mapping and monitoring koala habitat suitability for conservation management.

Aim The strength of consumer–plant interactions may decrease with latitude. Our objectives were to assess the spatial variation in folivory on Nothofagus pumilio and understand the influence of climate on folivory patterns as mediated by changes in folivore density and leaf traits. Location Nothofagus pumilio forests, between 38 and 55°S (Argentina). Methods We studied the correlation of leaf damage with latitude on data from 47 sampling sites, and evaluated spatial patterns of autocorrelation on latitudinally detrended data with a principal coordinates of neighbour matrices method. Path analysis was used to test the association of temperature and precipitation with leaf damage, mediated by folivore density and leaf traits. We evaluated the adequacy of this ecological model by examining the spatial pattern of autocorrelation in the residuals, and combined spatial and environmental predictors of leaf damage into partial regression. Results Leaf damage decreased with latitude, which was the only significant spatial predictor. The latitudinal decrease in temperature and precipitation was correlated with a decrease in the density of folivores and leaf size, and diminished leaf damage. Our ecological model adequately explained the spatial autocorrelation in the data: 44% of the variation in leaf damage was explained by the latitudinally structured component of the environment, whereas local environmental effects accounted for another 22%. Main conclusions We conclude that N. pumilio forests show consistent latitudinal patterns of variation in folivory, folivore density and leaf traits. Our study suggests that the latitudinal variation in folivory rates is partly driven by the influence of climate on both plants and herbivores. This warns us about the potential susceptibility of folivory rates to climate warming. We emphasize the value of large-scale analyses as complementary to local experimental approaches to understanding the regulation of herbivory.

Primate life histories are strongly influenced by both body and brain mass and are mediated by food availability and perhaps dietary adaptations. It has been suggested that folivorous primates mature and reproduce more slowly than frugivores due to lower basal metabolic rates as well as to greater degrees of arboreality, which can lower mortality and thus fecundity. However, the opposite has also been proposed: faster life histories in folivores due to a diet of abundant, protein-rich leaves. We compared two primate taxa often found in sympatry: Asian colobines (folivores, 11 species) and Asian macaques (frugivores, 12 species). We first described new data for a little-known colobine (Phayre's leaf monkeys, Trachypithecus phayrei crepusculus) from Phu Khieo Wildlife Sanctuary, Thailand. We then compared gestation periods, ages at first birth, and interbirth intervals in colobines and macaques. We predicted that heavier species would have slower life histories, provisioned populations would have faster life histories, and folivores would have slower life histories than frugivores. We calculated general regression models using log body mass, nutritional regime, and taxon as predictor variables. Body mass and nutritional regime had the predicted effects for all three traits. We found taxonomic differences only for gestation, which was significantly longer in colobines, supporting the idea of slower fetal growth (lower maternal energy) compared to macaques and/or advanced dental or gut development. Ages at first birth and interbirth intervals were similar between taxa, perhaps due to additional factors (e.g., allomothering, dispersal). Our results emphasize the need for additional data from wild populations and for establishing whether growth data for provisioned animals (folivores in particular) are representative of wild ones.

We studied the feeding ecology of the thin-spined porcupine (Chaetomys subspinosus), a small arboreal rodent endemic to the Atlantic rain forest of Brazil and threatened with extinction. We captured and radiotracked 4 free-ranging individuals and collected 1,177 feeding records in 944 h of observation between April 2005 and September 2006. We found that the animal feeds exclusively on the leaves of woody trees, preferring young leaves. Diet composition and foraging pattern did not vary seasonally, although we observed a seasonal variation in the availability of young leaves. Annual diet comprised primarily (90%) 4 plant species, Albizia pedicellaris, Inga thibaudiana, Pera glabrata, and Tapirira guianensis. This small subset of plant species contains high levels of both leaf protein and fiber, and most preferred species have the capacity to establish symbiotic associations with nitrogen-fixing bacteria. A diet concentrated in a few taxonomically related, nitrogen-fixing species suggests that in a...

The individual and combined effects of elevated CO 2 and 0 3 on the foliar chemistry of silver birch (Betula pendula Roth) and on the performance of five potential birch-defoliating insect herbivore species (two geometrid moths, one lymantrid moth and two weevils) were examined. Elevated CO 2 decreased the water concentration in both short- and long-shoot leaves, but the effect of CO 2 on the concentration of nitrogen and individual phenolic compounds was mediated by 0 3 treatment, tree genotype and leaf type. Elevated 0 3 increased the total carbon concentration only in short-shoot leaves. Bioassays showed that elevated CO 2 increased the food consumption rate of juvenile Epirrita autumnata and Rheumaptera hastata larvae fed with short-and long-shoot leaves in spring and mid-summer, respectively, but had no effect on the growth of larvae. The contribution of leaf quality variables to the observed CO 2 effects indicate that insect compensatory consumption may be related to leaf age. Elevated CO 2 increased the food preference of only two tested species: Phyllobius argentatus (CO 2 alone) and R. hastata (CO 2 combined with 0 3 ). The observed stimulus was dependent on tree genotype and the measured leaf quality variables explained only a portion of the stimulus. Elevated 0 3 decreased the growth of flush-feeding young E. autumnata larvae, irrespective of CO 2 concentration, apparently via reductions in general food quality. Therefore, the increasing tropospheric 0 3 concentration could pose a health risk for juvenile early-season birch folivores in future. In conclusion, the effects of elevated 0 3 were found to be detrimental to the performance of early-season insect herbivores in birch whereas elevated CO 2 had only minor effects on insect performance despite changes in food quality related foliar chemistry.

Petauroides volans (Kerr, 1792) is a pseudocherid marsupial with the common name greater glider. It is the largest and most conspicuous of the gliding possums and is 1 of the most numerous arboreal marsupials in the eucalypt forests of eastern Australia. Its gliding membranes are large and its thick, shaggy coat obscures the basic body form, making the animal appear much larger than it actually is. P. volans is nocturnal, arboreal, solitary, and folivorous and is dependent on large tree hollows for shelter. In Australia, P. volans has received particular attention because it is highly sensitive to the removal of old hollow-bearing trees during forestry practices such as clear-cutting.

Bradypus variegatus Schinz, 1825 (the brown-throated three-toed sloth), is 1 of 4 extant three-toed sloths. A high-canopy folivore, B. variegatus is distributed over southern Central America and northern and central South America and is listed as Least Concern by the International Union for Conservation of Nature and Natural Resources because of its wide distribution. Males have a middorsal speculum. B. variegatus has a commensal relationship with the algae and invertebrates that live in its thick pelage. It does not adapt well to zoo settings.

Because of the large differences between flower buds and leaves as food resources we predict that florivore caterpillars compared with folivores would present: (1) higher degree of dietary specificity; (2) higher abundance; and (3) lower species richness. Based on emerged adults from caterpillars collected on leaves and inflorescences our results showed that the florivore fauna was strongly dominated by a few species with higher abundance and dietary specificity than the folivore fauna.

Bradypus torquatus Illiger, 1811 is a three-toed sloth commonly called the maned three-toed sloth. It is endemic to the Atlantic coastal forests of southeastern Brazil. Occasionally placed in its own genus (Scaeopus), the black mane of this three-toed sloth is distinctive. Like other sloths, B. torquatus is a high-canopy folivore with a commensal relationship with algae and invertebrates that live in its abundant pelage. B. torquatus is an endangered species that does not adapt to captive zoo settings.

Plants contain a variety of chemical defenses that strongly affect feeding rates in captive mammals, but their effects on the fitness of wild herbivores are largely unknown. This is because the complexity of defensive compounds, and herbivores' counteradaptations to them, make their effects in the wild difficult to measure. We show how tannins interact with protein to produce spatial variation in the nutritional quality of eucalypt foliage, which is related to demography in a wild population of a marsupial folivore, the common brushtail possum (Trichosurus vulpecula Kerr). Tannins reduced the digestibility of nitrogen (N) in vitro, creating variation in available N concentrations among the home ranges of individual possums in an otherwise homogeneous habitat. This was strongly correlated with reproductive success: females with better quality trees in their home range reproduced more often and had faster-growing offspring. These results demonstrate a powerful mechanism by which spatial variation in plant chemistry may control herbivore population dynamics in nature.

Socioecological models indicate that the group structure and female dispersal patterns of primates are determined primarily by the abundance and distribution of food, predation pressures, and infanticide risks. In response to those influences, females of folivorous primates are considered relatively free to disperse into groups with the optimal size and structure. Yet some folivores live in small groups despite a potentially higher risk of predation, an apparent inconsistency known as the folivore paradox. This paper examines the female dispersal of a folivorous primate, the Virunga mountain gorillas. Mountain gorillas currently have no natural predators, but this species presents a different version of the folivore paradox: why do 50–60% of females reside in smaller one-male groups despite a higher risk of infanticide? In this study, females left one-male groups more frequently than multimale groups, but transfer destinations were not consistently biased toward multimale groups and those groups did not have higher immigration rates. We found no evidence of dispersal to avoid feeding competition within large groups, even as they have become three to five times larger than average. Thus, the lack of a consistent bias toward multimale groups was not because they are typically larger than one-male groups. Instead, the apparent inconsistencies may reflect limited female transfer opportunities, other influences on dispersal, and possibly an evolutionary disequilibrium in which current behavior does not optimize fitness.

Insect herbivores were collected from five species of dipterocarp tree seedling within a large-scale reciprocal transplant experiment in Sabah, Malaysia, on alluvial and sandstone soils in both gap and understory plots. The aim was to determine whether the location and ecological specialization of seedlings influenced the herbivore communities found on and around them. Three major groups of folivores were collected: Coleoptera, Orthoptera, and larval Lepidoptera. Herbivory of all species was confirmed through laboratory trials. Herbivore abundance in the understory plots was extremely low relative to the gaps. Rank-abundance curves were similar on both soil types, differing only within the Lepidoptera. Coleoptera and Orthoptera communities were numerically dominated by a small suite of species capable of feeding on all dipterocarp species tested, whereas lepidopteran communities had both greater species richness and diversity. When corrected for leaf area surveyed, the abundance of Coleoptera was similar on both soil types, while larval Lepidoptera were more abundant in sandstone plots and Orthoptera were more abundant in alluvial plots. Estimated species richness of all three taxa was greater in alluvial forest, but there were contrasting patterns in Simpson diversity and evenness between groups. Species richness of Lepidoptera was greatest on seedlings when grown in their native soil type, providing partial evidence for possible escape effects, although this was not matched by differences in folivore abundance. The link between herbivore communities and herbivory rates on rain forest tree seedlings is complex and is unlikely to be detected through simplistic measures of abundance, species richness, or diversity.

Numerous comparative studies have sought to demonstrate a functional link between feeding behavior, diet, and mandibular form in primates. In lieu of data on the material properties of foods ingested and masticated, many investigators have relied on qualitative dietary classifications such as "folivore" or "frugivore." Here we provide the first analysis of the relationship between jaw form, dietary profiles, and food material properties in large-bodied hominoids. We employed ratios of area moments of inertia and condylar area to estimate moments imposed on the mandible in order to evaluate and compare the relative ability to counter mandibular loads among central Bornean orangutans (Pongo pygmaeus wurmbii), Virunga mountain gorillas (Gorilla beringei beringei), and east African chimpanzees (Pan troglodytes schweinfurthii). We used data on elastic modulus (E) of fruit, fracture toughness (R) of fruit, leaves, and non-fruit, non-leaf vegetation, and derived fragmentation indices ( R/E and ER), as proxies for bite force. We generated bending and twisting moments (forcexmoment arm) for various mandibular loading behaviors using food material properties to estimate minimally required bite forces. Based on E and R of foods ingested and masticated, we hypothesized improved resistance to mandibular loads in Pongo p. wurmbii compared to the African apes, and in G. b. beringei compared to Pan t. schweinfurthii. Results reveal that our predictions are borne out only when bite forces are estimated from maximum R of non-fruit, non-leaf vegetation. For all other tissues and material properties results were contrary to our predictions. Importantly, as food material properties change, the moments imposed on the mandible change; this, in turn, alters the entire ratio of relative load resistance to moment. The net effect is that species appear over- or under-designed for the moments imposed on the mandible. Our hypothesis, therefore, is supported only if we accept that maximum R of these vegetative tissues represents the relevant mechanical property influencing the magnitude of neuromuscular activity, food fragmentation, and mandibular morphology. A general implication is that reliable estimates of average and maximum bite forces from food material properties require that the full range of tissues masticated be tested. Synthesizing data on ingestive and masticatory behaviors, the number of chewing cycles associated with a given food, and food mechanical properties, should inform the broader question of which foods and feeding behaviors are most influential on the mandibular loading environment.

The Hoatzin Opisthocomus hoazin is the only known bird with a well-developed foregut plant fermentation system; most fermentation takes place in the crop and caudal oesophagus. To test Hoatzin digestive efficiency, balance (total collection) trials with captive Hoatzins were made using two experimental diets of different composition and fibre content. Dry matter (DM) intakes were similar for the diets (mean = 62.8 g DM/kg body mass/ day). Average DM, organic matter and nitrogen digestibilities were not significantly different between diets, with average values of 72.9%, 75.0% and 78.3%, respectively. In vitro organic matter digestibilities by cow ruminal inoculum were very similar to organic matter digestibilities in live Hoatzins for both diets. Fibre digestibility was among the highest recorded for herbivorous birds. Cellulose and acid detergent fibre digestibilities were 58.8% and 52.7%, respectively. Neutral detergent fibre (NDF) digestibility differed among diets—the higher the NDF content of the diet, the higher the NDF digestibility. The NDF digestibilities were 37.9% and 70.9% for the two diets with NDF concentrations of 32.4% and 37.3%, respectively. Differences in NDF digestibility can be attributed to the different concentrations of hemicellulose in the experimental diets. The high overall digestibility by captive Hoatzins is higher than values previously reported for other avian herbivores but similar to those of foregut-fermenting mammals on similar diets. The unique digestive strategy of the Hoatzin maximizes digestion of cell wall and cell contents. The high digestive efficiency in the Hoatzin is not predicted by allometric models of fibre digestion as a function of body mass. Other nutritional benefits, such as detoxification of plant secondary compounds and microbial synthesis of essential amino acids and vitamins, may explain the evolution of foregut fermentation in this avian folivore.

The objective of this study was to determine the effects of gall induction on leaf phenolic compounds and their indirect effects on the subsequent attack of folivorous insects in Achatocarpus gracilis, Cordia alliodora, Guapira macrocarpa, Guettarda elliptica and Ruprechtia fusca that occur in both hillside and riparian sites at Chamela-Cuixmala tropical dry forest in western Mexico. There are differences in soil water content between riparian and hillside sites where trees in the first are mainly evergreen and deciduous in the second. A few tree species occur in both sites and their intraspecific phenological response is also different between sites. In this case, trees of a given species that occur in riparian sites will be evergreen whereas trees on hillsides of the same species will be deciduous. Four plant species had significantly greater total phenol concentrations in galled than ungalled leaves in both deciduous hillside and riparian sites. In three plant species associated with galls, host total phenol concentrations were significantly greater in short than in tall plants. The frequency and amount of folivore damage were greater in leaves without galls than leaves with galls in these four plant species. These results indicate that galling insect species may directly affect leaf phenolic concentrations and indirectly may affect the incidence and consumption of folivorous insects in tropical plant species. This may have important consequences on the preference of leaves by folivorous insects that might be excluded by galling insect species in this dry tropical system.

Plants interact with many different species throughout their life cycle. Recent work has shown that the ecological effects of multispecies interactions are often not predictable from studies of the component pairwise interactions. Little is known about how multispecies interactions affect the evolution of ecologically important traits. We tested the direct and interactive effects of inter- and intraspecific competition, as well as of two abundant herbivore species (a generalist folivore and a specialist aphid), on the selective value of a defensive chemical compound in Brassica nigra. We found that investment in chemical defense was favored in interspecific competition but disfavored in intraspecific competition and that this pattern of selection was dependent on the presence of both herbivores, suggesting that selection will depend on the rarity or commonness of these species. These results show that the selective value of ecologically important traits depends on the complicated web of interactions present in diverse natural communities and that fluctuations in community composition may maintain genetic variation in such traits.

Food competition is an expected cost of group living. It is therefore puzzling that there is little evidence for competition among group-living folivorous monkeys, for example, daily travel distance does not seem to increase with group size. It is even more puzzling that folivores do not form larger groups despite this apparent lack of food competition. This has become known as the folivore paradox, and to date, there is no broadly accepted theoretical solution. However, there have been no multigroup studies that have controlled for the potentially confounding effects of variation in habitat quality. We studied 9 groups of red colobus monkeys (Procolobus rufomitratus) in Kibale National Park, Uganda, and controlled for spatial and temporal variation in food availability. We found that larger groups occupied larger home ranges than smaller groups and that group size was related to increased foraging effort (longer daily travel distance), increased group spread, and reduced female reproductive success. Our results also suggest that monkeys in larger groups spent more time feeding and less time engaged in social behavior. These results suggest that folivorous red colobus monkeys experience within-group scramble competition and possess a suite of behavioral responses that may mitigate the cost of competition and represent adaptations for group living. The results offer insight into the folivore paradox and the evolutionary ecology of group size. Copyright 2008, Oxford University Press.

Although koalas are iconic Australian animals, no pharmacokinetic studies of any first-line medicines used to treat diseased or injured koalas had been published prior to 2010. Traditionally, medicine dosages suggested for this species underwent linear extrapolation from those recommended for domesticated species. The koala, a specialist folivore whose natural diet consists of almost exclusively Eucalyptus spp. foliage has anatomical and physiological adaptations for detoxifying their diet which also affect medicine pharmacokinetic profiles. This review addresses aspects of medicine absorption, clearance, and other indices (such as medicine binding to plasma proteins) of enrofloxacin/marbofloxacin and chloramphenicol used for the systemic treatment of chlamydiosis, and fluconazole ± amphotericin, and posaconazole for the treatment of cryptococcosis. Based on observations from published studies, this review includes suggestions to improve therapeutic outcomes when administering medicines to diseased koalas.

Herbivory, the feeding on living plant parts by animals, is a key ecosystem process that has widely recognized effects on primary production and on vegetation structure and composition. The effect of herbivory depends on herbivore feeding type and intensity. A number of methods have been used to measure the intensity and effects of herbivory. Herbivory generally is lowest in some forests and highest in aquatic ecosystems. Insects are the primary herbivores in forest ecosystems and may account for the bulk of herbivory in grasslands, even though vertebrate grazers are more conspicuous. Herbivory has well-known effects on survival, productivity, and growth form of individual plants. However, the traditional view of herbivory as a negative effect on plants is being replaced by a view that recognizes more complex effects of variable intensity and timing. Moderate intensities of herbivory often stimulate production, through compensatory growth, and flowering, thereby increasing fitness. Herbivores affect the turnover of plant nutrients to litter as plant fragments, feces animal tissues and nutrients leached from chewed surfaces. Herbivory also can affect climate and the likelihood and intensity of future disturbances.

In multi-predator systems with different predator ‘archetypes’, prey have an advantage when applying ‘archetype’-specific responses. However, antipredator responses can be lost or modified during times of ‘relaxed predation selection’. As a consequence, prey may become vulnerable when novel predators are introduced into their environments. This study investigates whether the Lumholtz’s tree-kangaroo (LTK) (Dendrolagus lumholtzi), an arboreal folivore marsupial of rainforests in North Queensland, Australia, (1) recognizes odor cues from arboreal and terrestrial predators, (2) exhibits archetype-specific antipredator responses to them, and (3) is able to transmit archetype-specific antipredator responses to novel predators of similar archetype. Five captive LTKs were exposed to lavender oil as neutral odor, fecal material from an extant arboreal predator, the amethystine python (Morelia amethystine), from two novel terrestrial predators, the domestic dog (Canis lupus familiaris) and the dingo (Canis lupus dingo), and a control odor (water). All subjects showed an increase in durations and frequencies of vigilance, and a decrease in the duration of comfort behaviors when exposed to the fecal material compared to the control odor, but not when the neutral odor was presented. This suggests that the presented fecal material is associated with predatory threats. Subjects increased duration and frequency of their movements in trials with fecal material from a python and a dingo, but not in trials with fecal material from a dog or the neutral odor which indicates that tree-kangaroos do not respond differently to arboreal and terrestrial predators. This conclusion is discussed with respect to the relatively recent descent of tree-kangaroos from rock-wallabies at a time when its ancient terrestrial predators became extinct, and with respect to reported fatalities of tree-kangaroos when encountering domestic dogs or dingoes.

1. Small mammalian folivores, such as the koala, are considered to be energetically limited by their relatively small gut capacity compared with metabolic requirements and the gut-filling effect of their poorly digestible leaf diet. However, during peak lactation, female koalas increase their food intake (35%) to meet the nutritional demands of reproduction. 2. This study examines mechanisms by which reproductive female koalas overcome limitations to food intake. 3. Digesta retention in the gastrointestinal tract was measured in free-living koalas using inert solute (cobalt (Co) complexed with EDTA) and particle markers (chromium (Cr)-mordanted to cell wall constituents' 600–1180 μm in diameter). 4. The whole gut rate of passage of both markers was 2–3 times faster than in captive koalas, probably because of 35%–69% greater food intakes in the wild. As in captive koalas, the solute marker was retained longer than the particle marker in free-living animals, indicating selective retention of fluid, solutes and small particles (including bacteria) in the hindgut caecum and proximal colon. The digesta retention of both markers were unaffected by the 35% increase in food intake associated with reproduction, largely due to a 42% increase in the solute marker pool size in lactating animals. The pool size of large particles in the digesta was unchanged. 5. Thus, female koalas meet the nutritional demands of reproduction at least partly by an increased solute digesta pool size, minimizing the detrimental effects of increased food intake on digestion of solutes and small particles, and on faecal loss of microbial protein. There was some indication that they also increase the passage rate of large particles or increase the efficiency of separation of large and small particles to reduce the gut-filling effects of large, poorly digestible particles. 6. Clearly the digestive strategy of the koala is more flexible, and limitations to food intake less stringent, than previously thought. Female koalas accommodate the increased food intake required to meet the demands of free-living and reproduction without compromising nutrient extraction from their eucalypt leaf diet. We suggest that similar flexibility in digestive strategy is likely to play an important role in the way that most small mammalian herbivores, especially arboreal folivores, meet the nutritional demands of reproduction.

We examined the spatial and temporal distribution of the foods of ursine colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana as a means to predict the monopolizablity and usurpability of their food resources. Recent evidence suggests that food may not be limiting for folivorous primates, and that male sexual coercion may be a more important influence on folivore social organization. To address the question, we collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000-August 2001) on 2 groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also conducted phenological monitoring and a tree survey of the two-group home ranges to establish food availability and distribution. We used 2 behavioral or organism-defined indicators of feeding behavior to assess potential resource contestability: food site residence time and distance moved between food sites. The colobus fed on a high diversity of species, most of their food trees were not clumped in distribution, within-tree interfood distances were short, and food trees were large. The only condition associated with the potential for monopolization was low food tree density. However, low food tree density may be offset by the colobus’ use of large trees. Taken together, the ecological and behavioral indicators suggest the food resources of Colobus vellerosus had a low potential for monopolization. Our results also indicate mature leaves had the longest food site residence time, which may suggest they should be the most usurpable plant part, though their presumed low quality and high abundance probably counteracted the effect. The pattern implied the potential for direct feeding competition among Colobus vellerosus at Boabeng-Fiema was low and agonistic interactions over food are not expected. Instead, a group size effect on feeding efficiency should be a more predominant influence on feeding efficiency, if food is limiting for the species.

We examined the effects of light availability on the suitability of foliage from red maple (Acer rubrum L.) (Aceraceae), black cherry (Prunus serotina Ehrhart) (Rosaceae), and sassafras [Sassafras albidum (Nuttall) Nees (Lauraceae)] for the larvae of the promethea moth, Callosamia promethea Drury (Lepidoptera: Saturniidae), in midseason. We identified replicate sun- and shade-grown saplings of red maple, black cherry, and sassafras from naturally growing populations in the field. Foliage collected from the experimental saplings was bioassayed using early instars of the promethea moth and assayed for nitrogen and carbon content. Promethea moth-larval performance and survivorship was highest on sassafras, intermediate on black cherry, and lowest on red maple. Larvae feeding on foliage from plants grown in the sun performed better than from those grown in the shade; the effect of light did not depend on sapling species. Foliar nitrogen content varied among the sapling species and was higher, overall, in foliage from plants grown in the sun. Nitrogen concentration related strongly and positively with larval performance and accounted for a great deal of the variation in performance both among the sapling species and between the sun and shade treatments. During midseason, foliar nitrogen content is determined by light availability, it varies among sapling species, and it is likely the primary constituent determining host quality for folivores on these sapling species.