The nutritional value of feeding on crops: Diets of vervet monkeys in a humanized landscape

Anthropogenic influences have dramatically altered the environments with which primates interact. In particular, the introduction of anthropogenic food sources to primate groups has implications for feeding behaviour, social behaviour, activity budgets, demography and life history. While the incorporation of anthropogenic foods can be beneficial to primates in a variety of nutritional ways including increased energetic return, they also carry risks associated with proximity to humans, such as risk of being hunted, disease risk and risk of conflict. Given such risks, we initiated a 3‐year study where we sought to understand the underlying nutritional motivations for anthropogenic food resource use by vervet monkeys (Cercopithecus aethiops) in the humanized matrix surrounding the Nabugabo Field Station in central Uganda. Feeding effort, defined as proportion of feeding scans spent on anthropogenic food, was not associated with ripe fruit availability nor with crop availability as determined by phenological monitoring. Likewise, there was no difference in the protein, fibre, or lipid composition of crop food items compared to wild food items. Individuals spent less time feeding overall in months over the 3 years with a higher proportion of time spent feeding on crop foods, suggesting a potential benefit in terms of accessibility (reduction in the proportion of activity budget devoted to feeding).     

The feeding ecology of Stephen's lory and nectar availability in its food plants

Stephen's lory, Vini stepheni , is one of four landbirds which are endemic to Henderson Island and until recently, its habits and ecology were unknown. I studied the diet and feeding ecology of Stephen's lory with particular reference to nectar variation in dominant food plants. Stephen's lory feeds on nectar, pollen and fruits from a variety of plants with lepidopteran larvae also forming a large component of the diet. The two plants which were dominant nectar sources for Stephen's lory produced flowers with very different nectar yields. Scaevola sericea flowers contained larger maximum volumes of more concentrated nectar than Timonius polygamus flowers. However, Scaevola flowers were a less predictable nectar resource, and grew at much lower densities, than Timonius flowers. Lories fed on the two flower types at different frequencies and it was hypothesized that the net rate of energy gained from feeding from a similar sized patch of the two plant species was comparable. Although nectar volumes of Timonius polygamus flowers decreased from the early mornings up until late afternoons, the lories did not show a corresponding daily pattern of foraging behaviour. Nectar yields also increased with the age of Timonius flowers. Stephen's lory has probably been the least affected of its genus by adverse changes to its habitat, and any information gained on the species may be useful when conservation measures of other Vini species are addressed.     

Switching to a feeding method that obstructs vision increases head-up vigilance in dabbling ducks

A major assumption in most models of foraging is that feeding and vigilance are mutually exclusive. A recent experimental study challenged this hypothesis and demonstrated that birds are able to detect predators while pecking seeds on the ground (head-down vigilance). Experimental obstruction of head-down vigilance makes birds increase head-up vigilance (i.e. the classical overt vigilance posture). For many foragers in the wild, visibility varies between habitats and foraging methods. We compared the vigilance of Teal Anas crecca and Shoveler Anas clypeata when foraging with their eyes above the water surface (shallow feeding, only the bill submerged) and when foraging with their eyes underwater (deep feeding, head and neck underwater, or upending), at three wintering sites in western France. Birds of both species spent less time in head-up vigilance during shallow foraging than during deep foraging, with no significant difference between sites, which suggests that they are capable of some vigilance during shallow foraging. During deep foraging, the time spent vigilant increased because the frequency of scans was much higher, while scan length decreased. However, these differences could have resulted from variations in the availability of food at different depths. In an experiment where the food availability was constant, we observed the same pattern, with a higher frequency of scans during deep foraging. This study therefore provides strong support for the idea that vigilance and foraging are not always mutually exclusive and shows that switching between searching methods can cause vigilance time – and, as a consequence, loss of feeding time – to vary. This should be taken into account in future field and experimental studies of the trade-off animals make between vigilance and feeding.     

Time budget of the green monkey,Cercopithecus sabaeus: Some optimal strategies

Members of a single group of green monkeys spent, on average, 44.8% of their waking time foraging, 46.7% resting, and 8.5% in social activities, over 1 year. There was significant variation in activity budgets over months (ranging from 35 to 55% of the time spent feeding). Diurnal rhythms of feeding and ranging were influenced by the daily cycle of temperature in predictable ways in different seasons: in the dry season, activity was reduced if it was too hot or too cold, while the temperature in the wet season did not affect activities. Feeding was also synchronized among individuals on a finer time scale, irrespective of the time of day. There was closer synchrony when feeding on less common foods. An ecological model of foraging time and energetics was tested, using estimates of the costs and benefits of foraging and predicting how these are optimally balanced in relation to the food density. Both feeding time and distance traveled increased as food availability increased. Costs and benefits were balanced over several days. Comparisons between populations of Cercopithecus aethiopswere made; differences in time budgets were compared with differences in the availability and quality of food. Insufficient comparative data are available for firm conclusions about the role of different energetic and nutritive strategies in population differences.     

Enclosure environment affects the activity budgets of captive Japanese macaques (Macaca fuscata)

Individuals adapt to changes in their environment, such as food availability and temperature, by adjusting the amount of time spent in different behavioral activities. These adjustments in behavior should vary across age-sex class according to specific physiological and social needs. We studied the activity budgets of three social Japanese macaque groups inhabiting either vegetated or nonvegetated enclosures in order to compare the effects of access with vegetation, as both food and substrate on resting, feeding, grooming and moving activities over a 12-month period. Daily access to natural foods for monkeys in the vegetated enclosure seems to be largely responsible for the differences in daily time budgets of these three groups. Resting time in all three groups was longer than the time devoted to other activities. Resting and moving time in the two nonvegetated enclosures was significantly longer than in the vegetated enclosure. In contrast, feeding and grooming time was significantly longer in the vegetated enclosure. Seasonal variation in time spent feeding, resting and grooming was significantly effected by enclosure type. In all three enclosures, immatures, particularly females, spent more time feeding and moving, whereas adults spent more time resting. Significant monthly variation in time spent by age-sex class was noted only for feeding and resting. Interestingly, in the vegetated enclosure, time spent feeding on natural vegetation was equal to the amount of time spent feeding on provisioned food. This suggests that factors other than energetic and nutritional needs may be important determinants of the activity budget of the species. These results have important implications for the enrichment of captive primates and our understanding of the maintenance of activity patterns by primates in the wild.     

Time budgets of grassland herbivores: body size similarities

Summary The summer (May–September) time budgets of 14 generalist herbivore species living in the same grassland environment are presented in terms of various component activities (e.g., walking, feeding, resting, etc.). All the species exhibit a decrease in activity as average daily air temperature increases. Greater body size and variety of habitats used by a species lead to increased time spent active. Use of a greater variety of habitats may increase activity time because different habitats provide suitable thermal conditions for activity at different times of the day. Body size affects sn herbivore's thermal balance through metabolism, body surface area and thermal inertia. The time spent feeding, exclusive of time spent searching for foods, is less for large than small herbivores. This may arise because large species must spend more time walking in the search for food to satisfy their energy requirements. The observed feeding time differences for species composing a common trophic level in a single environment may help to explain their diet choice because feeding time constrains the variety of foods an herbivore can select. Diet differences, in turn, can explain the potential competition for food if food is in short supply.     

Garden bird feeding predicts the structure of urban avian assemblages

Households across the developed world cumulatively spend many millions of dollars annually on feeding garden birds. While beneficial effects on avian assemblages are frequently claimed, the relationships between levels of garden bird feeding and local avian populations are unknown. Using data from a large UK city, we show that both avian species richness and abundance vary across different socioeconomic neighbourhood types. We examined whether patterns in bird feeding could explain this variation. The density of bird feeding stations across the urban environment was strongly positively related to avian abundance, after controlling for differences in habitat availability. This effect was almost exclusively driven by the abundance of those species known to utilize garden feeding stations frequently. In contrast, the density of feeding stations had no effect on avian species richness. We also examined variation in the proportion of households in different communities that provide food for birds, a factor that is not correlated with feeder density. The prevalence of bird feeding across different neighbourhoods declined as socioeconomic deprivation increased, and increased with avian species richness and abundance. Our results suggest that the provision of supplementary food for birds by multiple landowners across a city can impact the status of urban bird populations. The potential for harnessing these actions for conservation needs to be explored.     

Resource base,parity, and reproductive condition affect females' feeding time and nutrient intake within and between groups of a baboon population

Summary We examined within- and between-group differences in aspects of feeding and nutrient intake among adult females of a single population of baboons (Papio cynocephalus) in Amboseli National Park, Kenya. Differences in time spent feeding, daily energy and protein intake and feeding efficiency (nutrient intake per minute spent feeding) reflected differences in resource base, reproductive condition and parity. Baboons that partially fed from a lodge garbage dump spent less than half the time feeding than those that were feeding totally in the wild. During this greatly reduced feeding time, the garbage-feeding group had a similar daily energy intake and only a slightly lower daily protein intake relative to wild-feeding baboons. Consequently, the feeding efficiency of the semi-provisioned baboons was appreciably higher than that of the non-provisioned baboons. For the totally wild-feeding baboons, samples were large enough to permit analyses of feeding time and nutrient intake during different reproductive states and parity. Females spent more time feeding and had higher daily energy and protein intake when they were pregnant or lactating than when they were sexually cycling. Nulliparous females spent more time feeding than their multiparous counterparts. The daily energy intake of nulliparous females was higher than that of their multiparous counterparts, but their daily protein intakes did not differ significantly. Pregnant or lactating and nulliparous females had higher feeding efficiency than their sexually cycling and multiparous counterparts. The two nulliparous females in the garbage-feeding group spent more time feeding but did not take in more energy or protein per day than their multiparous counterparts.     

Diet and feeding behaviour of samango monkeys (Cercopithecus mitis labiatus) in Ngoye Forest, South Africa

The samango monkey occurs at the southern limit of the range of Cercopithecus mitis. Greater climatic seasonality at this latitude results in more predictable fruiting patterns. In addition, there are no diurnal sympatric primate frugivores. Under these conditions, the diet and feeding strategies of samango monkeys would be expected to differ notably from those of central or east African C. mitis subspecies. Contrary to these expectations, the preliminary observations reported here indicate that diets of samango and blue monkeys differ only superficially in the proportions of items eaten. Similarities in feeding behaviour are especially marked during the dry season period when fruit is not abundant. Both samango and blue monkeys tend to be less selective in their choice of food species and to eat available food species regardless of their energy content; a shift toward less nutritious items such as leaves is also noted. Feeding behaviour during the summer wet season is characterized by the selection of fruits with high-energy values. A high proportion of visits by the monkeys to areas of greater food availability suggests a concentration of feeding effort in food patches and the selection of higher energy food species within patches.     

Seasonal Variations in the Diet and Feeding Effort of Two Groups of Howlers in Different Sized Forest Fragments

Little is known about how resource limitation affects the feeding ecology of primates in forest fragments. Here, we describe seasonal variation in the diet and feeding effort of 2 groups (RH and RC3) of howlers (Alouatta palliata mexicana) living in different sized forest fragments in Los Tuxtlas, Mexico. The RH group, which lived in a larger and more preserved forest fragment, with a higher availability of fruit and Ficus trees, had a higher and more constant consumption of fruit throughout the study year. Moreover, this group fed from larger food sources, i.e., trees, lianas, and shrubs, and spent more time feeding from each food source. The feeding effort, defined as the time spent feeding and traveling divided by time spent resting, of the RH group was also significantly lower and more stable than that of the RC3 group throughout the study year. As feeding effort has been positively related to stress in primates, such increases in feeding effort could have negative consequences for howlers in small or degraded forest fragments. Our study suggests that habitat characteristics interact with plant seasonality to determine the diet and feeding effort of howlers in forest fragments and that groups living in small and degraded forest fragments may be particularly vulnerable to years of low fruit production.     

The Ranging Costs of a Fallback Food: Liana Consumption Supplements Diet but Increases Foraging Effort in Howler Monkeys

Lianas are important components in the dynamics of tropical forests and represent fallback foods for some primates, yet little is known about their impact on primate ecology, behavior or fitness. Using 2 yr of field data, we investigated liana consumption and foraging effort in four groups of howler monkeys (two in bigger, more conserved forest fragments and two in smaller, less conserved fragments) to assess whether howler monkeys use lianas when and where food availability is scarce, and how liana consumption is related to foraging effort. Howler monkeys in smaller fragments spent more time consuming lianas and liana consumption was negatively related to the consumption of preferred food resources (fruit and Ficus spp.). Further, travel time was positively related to liana feeding time, but not to tree feeding time, and howler monkeys visited a greater number of food patches when feeding from liana leaves than when feeding from tree leaves. Our results suggest that these increases in foraging effort were related to the fact that lianas are mainly a source of leaves, and that liana patch size was probably smaller than tree patch size. While these results were clear when analyzing all four groups combined, however, they were not always significant in each of the groups individually. We suggest that this may be related to the differences in group size, patch size and the availability of resources among groups. Further studies are necessary to assess whether these dietary and behavioral adjustments negatively impact on the fitness and conservation of primates in fragments.     

Optimal feeding under stoichiometric constraints: a model of compensatory feeding with functional response

When the nutrient content of food is limited, herbivores often increase their feeding rates. Such an increase in the feeding rate is called ‘compensatory feeding’. Although it has a number of implications for herbivore population and plant–forager dynamics, the compensatory feeding is not yet functionally formulated especially in relation with ecological stoichiometry. Therefore, we constructed a simple mathematical model by incorporating the optimal feeding rate into the type II functional response to maximize a forager's growth rate under constraints of carbon or nutritionally important element like phosphorus (P). We used the planktonic herbivore Daphnia as a model herbivore. The model revealed that the optimal feeding rate increased by using excess carbon when relative P content of food was less than a certain level, which is known as the threshold elemental ratio. This level changed with the change of food abundance. It also showed that whether or not foragers should exhibit compensatory feeding depends on their stoichiometric characteristics and digestive traits, and also on the assimilability of a given food. These findings are helpful to test the feeding conditions under which compensatory feeding is advantageous for a given animal. Our model can be easily incorporated into forager population dynamics and prey‐consumer interaction models because the optimal feeding rate can be analytically given.     

Winter feeding of birds increases productivity in the subsequent breeding season

Supplementary food given to birds can have contemporary effects by reducing the risk of starvation, increasing survival and altering movements and reproductive performance. There is, however, a widely held perception that birds benefit from extra food over winter, but that it is better that they 'look after themselves' during breeding. Here we describe a landscape-scale experiment showing for the first time that the effects of increasing food availability only during the winter can be carried over to the subsequent breeding season. Even though food supplementation stopped six weeks prior to breeding, birds living on sites provisioned over winter had advanced laying dates and increased fledging success compared with birds living on unprovisioned sites. Thus, supplemental feeding of wild birds during winter, in a manner mimicking householders provisioning in gardens and backyards, has the potential to alter bird population dynamics by altering future reproductive performance. With levels of bird feeding by the public continuing to increase, the impacts of this additional food supply on wild bird populations may be considerable.     

Daily feeding rhythm in proboscis monkeys: a preliminary comparison with other non-human primates

In non-human primates, the daily feeding rhythm, i.e., temporal fluctuation in feeding activity across the day, has been described but has rarely received much analytical interpretation, though it may play a crucial part in understanding the adaptive significance of primate foraging strategies. This study is the first to describe the detailed daily feeding rhythm in proboscis monkeys (Nasalis larvatus) based on data collected from both riverbank and inland habitats. From May 2005 to May 2006, data on feeding behavior in a group of proboscis monkeys consisting of an alpha-male, six adult females and immatures was collected via continuous focal animal sampling technique in a forest along the Menanggul River, Sabah, Malaysia. In both the male and females, the highest peak of feeding activity was in the late afternoon at 15:00–17:00, i.e., shortly before sleeping. The differences in the feeding rhythm among the seasons appeared to reflect the time spent eating fruit and/or the availability of fruit; clearer feeding peaks were detected when the monkeys spent a relevant amount of time eating fruit, but no clear peak was detected when fruit eating was less frequent. The daily feeding rhythm was not strongly influenced by daily temperature fluctuations. When comparing the daily feeding rhythm of proboscis monkeys to that of other primates, one of the most common temporal patterns detected across primates was a feeding peak in the late afternoon, although it was impossible to demonstrate this statistically because of methodological differences among studies.     

Foraging behaviour and feeding locations of Rock Shags Phalacrocorax magellanicus from a colony in Patagonia, Argentina

During 1996 and 1997, foraging Rock Shags Phalacrocorax magellanicus were studied at Punta Loma, Argentina using radio-transmitters deployed on ten adult shags during the chick-rearing period. Rock Shags undertook 2.6 ± 0.6 sd trips per day. The mean duration of a feeding trip was 2.6 ± 0.7 hours. A bird spent 36% of daylight hours away from the colony on feeding trips, diving for 92% of the foraging trip, and made a mean of 106 dives per foraging trip. Foraging trip duration was strongly correlated with the total number of dives made in one foraging trip. Rock Shags fed mainly in water less than 10m deep with a gravelly sand bottom and within 5 km of shore. Mean foraging range was 3.8 ± 2.6 km and 2.6 ± 2.3 km for 1996 and 1997, respectively. These results suggest a high foraging effort (diving time per foraging trip) for Rock Shags, presumably associated with poor food conditions close to the colony. Comparison is made with other Phalacrocorax species.     

Behavioral responses of Cao Vit gibbon (Nomascus nasutus) to variations in food abundance and temperature in Bangliang, Jingxi, China

The Cao Vit gibbon is a critically endangered species with only about 110 individuals remaining in a degraded karst forest along the China-Vietnam border. Behavioral data from this site are particularly useful in understanding gibbon behavioral adaptations to different sets of ecological conditions and will contribute to the conservation of the species. We studied seasonal variation in the time budget and diet of the Cao Vit gibbon in response to variation in food availability and ambient temperature by observing two groups for 1,379 hr between January and December 2009. We used 5-min scan samples to record the activity of gibbons. Both ambient temperature and food availability varied from month to month. Gibbon groups increased resting time and huddled together in sleeping places in cold months. Gibbons spent more time feeding on fruit when fruit was more abundant suggesting that fruit was their preferred food. Alternatively, leaf eating was negatively correlated with leaf availability which suggested that leaves may be used as a fallback food. Gibbons increased their diet diversity when they ate more leaves. This might be a strategy to cope with toxins or digestion inhibitor accumulation associated with feeding from a limited number of leaf species. Individuals consumed more buds when Broussonetia papyrifera produced buds in March and April. During this period, they decreased traveling time and engaged in less frequent social interactions. Gibbons spent more time searching for and feeding on invertebrates during June and October. However, we did not collect data on invertebrate abundance and therefore cannot determine the relationship between invertebrate feeding and availability. We conclude that flexibility in consuming diverse food types and food species, and in responding to the availability of preferred foods, has enabled the Cao Vit gibbon to survive in a degraded karst forest habitat.     

Seasonal changes in food resource distribution and feeding sites selected by Japanese macaques on Koshima Islet,Japan

Feeding sites of Japanese macaques (Macaca fuscata) change according to seasonal fluctuations in food resource distribution. To examine what characteristics of food items affect feeding site selection, I describe herein the seasonal changes in food items, feeding sites, and food resource distributions of this species. Feeding behavior of monkeys and their food resource distributions were investigated on Koshima Islet, southern Japan, for four study periods (i.e., seasons) in 2002. Monkeys showed large variations in their diet between seasons. To weigh the relative influence of the distribution and abundance of food items on feeding site selection in each season, multiple regression analyses were performed by 100 m × 100 m grid. In the analyses, feeding time was a dependent variable and the abundance of staple food items, for which feeding time was over 5% in each season, in each grid square was an independent variable. There was no correlation between the resource distribution of most food items and the distribution of feeding time by monkeys in each season. Monkeys spent more feeding time where multiple staple food items were available. Food items that affected feeding site selection by monkeys had the following three characteristics: (1) clumped distribution, (2) seasonal availability, and (3) fruit. This suggests that monkeys are likely to select feeding sites to consume food items whose availability is limited temporally and spatially, which may enable them to simultaneously use other widely distributed, abundant food items efficiently.     

Choosing where to feed: the influence of competition on feeding behaviour of cod, Gadus morhua L.

Groups of cod, Gadus morhua L, presented with two feeding patches with a food abundance ratio of 2:1, distributed themselves between the patches in a ratio of 2.5:1. This is slightly higher than the 2:1 ratio predicted by the ideal free distribution theory. Large differences were observed in competitive ability between individual fish. A strong correlation was found between feeding success of individuals and time spent in a feeding patch. The more successful competitors caught about 2.5 times as many food items in the rich patch as in the poor patch. The less successful competitors caught an equal amount of food in both patches. All competitors, however, spent significantly more time in the rich patch. These results suggest that hunting success is the most important factor in assessing patch quality. However, it is not the only parameter which cod use in deciding where to feed.     

Adaptive regulation of body reserves in reindeer, Rangifer tarandus: a feeding experiment

The costs and benefits of body reserves fluctuate according to predictable factors such as season and life-cycle stage. Theory suggests that individuals at any time should regulate their body reserves according to the current balance between costs and benefits. Most studies on adaptive body mass regulation have been done on small passerine birds. In large vertebrates the costs associated with body reserves are assumed to be small and the reserves of these species are therefore thought to be dictated by environmental limitations. In this study we present experimental evidence for adaptive body mass regulation in female semi-domesticated reindeer ( Rangifer t. tarandus ). The risk of starvation in this species is highest in late winter. During snow melt this risk is reduced and the females should direct their effort towards the protection of their new born calf. To test how these seasonal and life-cycle changes are related to body mass regulation, we conducted a crossed experiment with two treatments where females were fed ad libitum during winter and spring respectively. During winter, the females from the fed group gained on average 12% of their initial body weight while the females from natural pastures lost on average 6% of their initial body weight. This strong response to winter feeding had no effect on reproductive performance, and the previously fed females lost their excess of body reserves during feeding in spring. This suggests that body reserves during winter primarily is used as an insurance against stochastic periods of starvation and that the females regulate their body reserves down to a set point in spring when the risk is reduced. We found however a positive correlation between initial female body weight and reproductive performance suggesting a close relationship between body weight and intrinsic individual qualities.     

Influence of food availability and climate on behavior patterns of western black crested gibbons (Nomascus concolor) at Mt. Wuliang,Yunnan, China

Food abundance and climatic factors can significantly affect the behavior of animals and constrain their activity budgets. The population of western black crested gibbons (Nomascus concolor) in Mt. Wuliang lives in montane forest and is close to the northern extreme of the distribution for gibbons (Hylobatidae). Their habitats show remarkable seasonal variation in terms of food availability, temperature, and rainfall. To understand behavioral adaptations of western black crested gibbons to different sets of ecological conditions, we examined relationships among food availability, mean temperature, rainfall, and behavior patterns by observing two groups for 1 year each. Our results revealed that activity budget was affected by food availability and mean temperature. The gibbons spent more time eating flowers when that resource was more available and spent less time moving when fruit was more available. The gibbons spent less time feeding and more time resting, and spent less feeding time on fruit and leaves when the mean temperature was lower. These results suggest that the gibbons displayed a pronounced preference for flowers as a food resource and adopted a time minimizer strategy when high‐nutrient food items (i.e., fruit) were more available. In addition, the gibbons adopted an energy‐conserving strategy during periods of low temperature. The flexibility of behavioral patterns in responding to food availability and temperature may potentially improve the gibbons’ prospects of surviving and reproducing in a northern montane forest.