Optimal foraging on the roof of the world: Himalayan langurs and the classical prey model

Optimal foraging theory has only been sporadically applied to nonhuman primates. The classical prey model, modified for patch choice, predicts a sliding “profitability threshold” for dropping patch types from the diet, preference for profitable foods, dietary niche breadth reduction as encounter rates increase, and that exploitation of a patch type is unrelated to its own abundance. We present results from a 1‐year study testing these predictions with Himalayan langurs (Semnopithecus entellus) at Langtang National Park, Nepal. Behavioral data included continuous recording of feeding bouts and between‐patch travel times. Encounter rates were estimated for 55 food types, which were analyzed for crude protein, lipid, free simple sugar, and fibers. Patch types were entered into the prey model algorithm for eight seasonal time periods and differing age‐sex classes and nutritional currencies. Although the model consistently underestimated diet breadth, the majority of nonpredicted patch types represented rare foods. Profitability was positively related to annual/seasonal dietary contribution by organic matter estimates, whereas time estimates provided weaker relationships. Patch types utilized did not decrease with increasing encounter rates involving profitable foods, although low‐ranking foods available year‐round were taken predominantly when high‐ranking foods were scarce. High‐ranking foods were taken in close relation to encounter rates, while low‐ranking foods were not. The utilization of an energetic currency generally resulted in closest conformation to model predictions, and it performed best when assumptions were most closely approximated. These results suggest that even simple models from foraging theory can provide a useful framework for the study of primate feeding behavior. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Shifting forest composition and primate diets: a 13-year comparison of the Tana River mangabey and its habitat

We compared the feeding behavior of a group of Tana River mangabeys (Cercocebus galeritus), densities of 25 diet species, and fruit availability of nine species in a 16.25 ha Tana River forest in southeastern Kenya studied in 1988 and in 2000-2001. For both studies, we enumerated all reproductively sized individuals of the 25 diet species and sampled nine of those species monthly for fruit availability. Mangabey feeding data were collected monthly from January to December 1988 and August 2000 to July 2001 using identical methods. We found a 17% increase in stem number of the 25 species between studies. Estimates of fruit production were lower in 2000 for five of the nine species monitored. Species composition of the mangabey diets shifted between 1988 and 2000-2001. We suggest that changes in forest composition may be due to declining human disturbance, elephant loss, changes in the river's hydrologic regime, and the 1997/98 ENSO event. Possible reasons for lower fruit availability are a younger demographic profile of the forest and changes in the river's hydrological regime. Only some of the changes in the mangabey diet mirrored changes in stem abundance and/or fruit availability. Mangabey dietary changes underscore their high degree of flexibility that allows them to persist in such a dynamic forest habitat.     

Predicting folivorous primate abundance: validation of a nutritional model

Understanding the determinants of animal abundance has become more vital as ecologists are increasingly asked to apply their knowledge to the construction of informed management plans. However, there are few general models are available to explain variation in abundance. Some notable exceptions are studies of folivorous primates, in which the protein-to-fiber ratio of foods has been shown to predict biomass. Here we examine the generality of Milton's [American Naturalist 114:363-378, 1979] protein/fiber model by providing a detailed analysis of diet selection in black-and-white colobus monkeys (Colobus guereza), and applying the model to populations shown to be stable; an assumption not previously examined. Based on observations of two groups of black-and-white colobus in Kibale National Park, Uganda, and one group in a forest fragment, we documented that the animals selected young leaves that had more protein, were more digestible, and had a higher protein-to-fiber ratio than mature leaves. The mature leaves did not differ from young leaves with respect to secondary compounds or mineral content (with the exceptions of copper and zinc). All of the colobus groups selected foods with a high protein-to-fiber ratios. However, one group also selected more digestible foods, and in another group, foraging efforts were positively related to zinc and negatively related to potassium. Previous studies that examined Milton's protein/fiber model did not demonstrate that the study populations were stable. If some populations were not at carrying capacity, then the correlations drawn between food availability and/or quality and folivore biomass may have been spurious. To address this issue, we censused a series of forest fragments in 1995 and again in 2000. We found that the populations in these fragments had declined from 165 in 1995 to 119 animals in 2000. However, based on evidence of population stability and lack of forest disturbance, we concluded that five of the original populations were stable. The biomass of these populations was related to the protein-to-fiber ratio of the fragment's trees. Combining our data with published data, we demonstrate that the protein-to-fiber ratios of mature leaves available to these folivorous primates accounted for 87% of the variance in their biomass.     

Diet ofColobus polykomos on Tiwai Island: Selection of food in relation to its seasonal abundance and nutritional quality

A group of Colobus polykomosat Tiwai, Sierra Leone, demonstrated seasonal flexibility in its diet, with seeds, young leaves, and mature leaves each dominating the diet at different times. Comparison of food consumption with phenological data indicates that seeds are eaten whenever available and are preferred to other foods, while young leaves are preferred to mature leaves. Colobus polykomosalso prefer liane to tree leaves, despite the relatively high quality of mature tree foliage at the Tiwai site. Analysis of protein, fiber, and energy values of foods selected and items available, but not eaten, suggests that preference is related to protein and energy maximization. Leguminous plants, especially Papilionaceae and Mimosaceae, are highlighted as important food sources for C. polykomos;seeds and leaves from these families have a high nitrogen content, and the protein content of leguminous seeds often equals or exceeds that found in leaves. It is predicted, therefore, that colobines living in habitats with a high density of legumes will feed heavily on seeds, subject to constraints such as seasonal availability. Where suitable leguminous species are less common, a mixture of fruits, seeds, and young or mature leaves or both is likely to be selected. The results of this and other recent studies of colobines do not support the notion that colobines are specialist folivores.     

The primate community of the Lopé reserve,Gabon: Diets,responses to fruit scarcity,and effects on biomass

The diets of all diurnal primates (Gorilla g. gorilla, Pan t. troglodytes, Mandrillus sphinx, Colobus satanas, Cercocebus albigena, Cercopithecus nictitans, C. pogonias, C. cephus) in the Lopé Reserve, central Gabon, are described from qualitative and quantitative data collected over 10 years. A total of 397 foods were recorded, of which 91% were from plants. The diets of seven of the eight species were numerically dominated by fruit, the exception being Colobus satanas with a diet dominated by seeds. When proportion of feeding time was examined, fruit remained the dominant food class for six species, while Cercocebus albigena, like the black colobus, spent most time feeding on seeds. The number of foods recorded per species ranged from 46–220, but dietary breadth of the four species that have not been subjects of detailed study was certainly underestimated. Almost half of the foods (46%) were eaten by only one species, and only four (1%) were recorded for all eight species. At Lopé, fruit is scarce for 2–3 months during the annual dry season, and fruit crop failures can prolong this period of scarcity to as many as 8 months in some years. When fruit is scarce, the diets of all species changed to include more nonfruit foods, but chimpanzees and the three species of guenons maintained a fruit dominated diet. The total biomass of the Lopé primate community is 374 kg/km², which is low compared to other sites in tropical Africa. Within the Lopé community, biomass correlates negatively with degree of frugivory, suggesting that fruit availability may be critical, but this does not explain the low total biomass compared to other sites. Neither floristic diversity nor the composition of the primate community at Lopé can account for the low biomass. While historical events such as climate-induced changes in forest cover, disease, or impact of human activities cannot be ruled out, we conclude that the most likely explanation of low primate biomass is the occurrence of an ecological “catastrophe” in the fairly recent past from which populations have not yet recovered. We believe that the most likely scenario was climate-mediated disturbance of flowering of important fruit food species. Prolonged fruit scarcity would have caused mortalities and affected the most frugivorous species more severely. Am. J. Primatol. 42:1–24, 1997. © 1997 Wiley-Liss, Inc.     

Food choice in Siganus fuscescens: influence of macrophyte nutrient content and availability

Using multiple‐choice feeding experiments, the selection of six species of macrophytes by the herbivorous rabbitfish Siganus fuscescens was examined. The rabbitfish showed distinct food choice in the laboratory; however, selection of macrophytes by S. fuscescens was not related to their absolute nutrient content (nitrogen, carbon, energy and ash free dry mass). Nutrient assimilation estimates showed that the macrophytes which were most preferred were those that S. fuscescens assimilated best. In S. fuscescens , the macrophytes that were preferred passed through the gut significantly faster than the less preferred species. Gut transit time had a significant effect on the absolute value of a food item in terms of net nutrient gain per unit time. This study showed that food value could be inferred from the absolute nutrient content of the macrophytes. Thus both the ability to assimilate nutrients as well as the absolute nutrient content of macrophytes must be quantified when assessing food value.     

Compensatory increases in food consumption and utilization efficiencies by velvetbean caterpillars mitigate impact of diluted diets on growth

Fresh weight (fw) food consumption by caterpillars of Anticarsia gemmatalis Hübner (Lepidoptera: Noctuidae) increased almost 2-fold as the nutrients in an artificial diet were increasingly diluted with water (diets contained 65, 79, 86 or 89% fw water). Nonetheless, dry weight (dw) relative consumption rate (RCR) declined with diet dilution. The efficiency at which the consumed food is digested and assimilated (approximate digestibility, AD) increased on the 3 diluted diets, and the efficiency at which digested food is converted to biomass (ECD) increased on the 79 and 86% fw diets. As a consequence, dw gained and relative growth rate (RGR), which is the product of RCR x AD x ECD, on the 79% fw diet were similar to those on the 65% fw diet, but they declined on the more diluted diets. Relative nitrogen consumption rate also declined with dietary dilution, but this was compensated by an increase in nitrogen utilization efficiency such that the product of these, relative nitrogen accumulation rate, was similar on all four diets. Insect lipid content declined from 32% on the undiluted diet to 13% dw on the most diluted diet, and was primarily responsible for the decline in RGR. The increases in fw consumption and AD, while not preventing a decline in RGR on the two most diluted diets, mitigated the impact of dietary dilution (e.g., without these increases, RGR on the most diluted diet would have been only 43% of that attained). These results indicate that the consumption and utilization of food are dynamic processes, and that caterpillars of A. gemmatalis, like many other insects, exhibit compensatory responses to changes in dietary quality.

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Résumé Peu de travaux ont examiné l'aptitude des insectes à modifier leur consommation et l'utilisation des aliments en fonction des variations qualitatives et quantitatives connues de l'aliment. On a rarement examiné si ces modifications sont adaptatives, si elles maintiennent le taux de croissance et le niveau des ressources, ou si elles limitent au moins les effets nuisibles du changement d'aliment. En utilisant une technique gravimétrique, nous avons mesuré le gain des poids, l'aliment consommé et les excréments produits par des chenilles de A. gemmatalis Hübn. (Lep, Noctuidea), important ravageur du soja aux USA.La consommation en poids frais (fw) a augmenté presque 2 fois quand le régime artificiel a été dilué progressivement avec de l'eau (65, 79, 86 ou 89% d'eau); le poids sec (dw) et le taux de consommation relative (RCR) ont diminué néanmoins avec la dilution. L'efficacité de digestion et d'assimilation de l'aliment consommé (digestibilité approchée, AD) a augmenté pour les trois dilutions, l'efficacité de conversion en biomasse de l'aliment digéré (ECD) a augmenté dans les dilutions à 79 et 85%, mais a diminué pour les régimes plus dilués. Par conséquent, le gain dw et le RGR, — produit de RCR x AD x ECD-, étaient identiques avec le régime à 79% fw à ceux de 65% fw, mais ont diminué pour les régimes plus dilués. Le taux relatif d'efficacité de consommation d'azote a diminué aussi avec la dilution, mais il a été compensé par un accroissement de l'efficacité de l'utilisation de l'azole tel, que le produit,-taux relatif d'accumulation de l'azote-, était le même pour les quatre régimes. La teneur en lipides des insectes a diminué, de 32% pour le régime sans dilution, à 13% pour le régime le plus dilué; elle a été le principal responsable de la diminution de RGR. L'accroissement de la consommation fw et de AD, tout en n'empêchant pas la diminution de RGR pour les deux régimes les plus dilués, a limité l'effet de la dilution (sans ces accroissements, le RGR du régime le plus dilué n'aurait été que 43% du RGR obtenu).Ces résultats indiquent que la consommation et l'utilisation d'aliments constituent un processus dynamique, et que les chenilles de A. gemmatalis comme beaucoup d'autres insectes, présentent des réactions compensatrices au changement de qualité de l'aliment. De telles variations dans la consommation d'aliments ont des conséquences écologiques (consommation accrue de substances allélochimiques potentiellement toxiques et exposition accrue aux causes biologiques de mortalité pendant l'alimentation), et intéressent l'organisation de la protection contre les insectes:-en permettant la manipulation de l'alimentation des insectes pour augmenter la consommation à court terme d'agents bioactifs, et, par ce moyen, réduire la consommation totale,-en permettant d'améliorer la modélisation des dégâts alimentaires prévisibles, qui, pour être efficace, devrait inclure les variations de l'alimentation en fonction de la qualité alimentaire de la plante cultiveé.

     

Sexual development of wild and mass‐reared male Queensland fruit flies in response to natural food sources

Diet has a profound influence on the fitness of adult tephritid flies. Mass‐reared flies are provided yeast hydrolysate as a rich source of nutrition that supports rapid sexual development and mating success. In contrast, wild tephritid flies often live in environments where food may be hard to find, and these are the conditions that sexually immature mass‐reared sterile males encounter when released into the field during sterile insect technique campaigns. The effect of natural food sources (bat guano, bird droppings, citrus pollen, and wheat pollen) on the sexual development of adult mass‐reared fertile, mass‐reared sterile, and wild male Queensland fruit flies, Bactrocera tryoni (Froggatt) (Diptera: Tephritidae), was determined by measuring ejaculatory apodeme size. Inclusion of yeast hydrolysate in the adult diet was associated with faster growth of the ejaculatory apodeme in comparison with all other diets. Effects of diet were far less pronounced in mass‐reared males, which may indicate reduced nutritional requirements, whereas the ejaculatory apodeme of wild males fed on natural sources of food or sucrose alone did not increase in size over the first 20 days of adult life.     

不同生态恢复措施对草地样带食物消费的影响——以锡林郭勒盟为例

生态工程的实施对区域居民生产生活产生了重要影响。以锡林郭勒草地样带为例,利用2017年8月—2018年7月农牧户食物跟踪调查数据、农牧户问卷调查数据、生态恢复措施数据,结合土地利用数据和社会经济数据等,重点分析了不同生态恢复措施对锡林郭勒草地样带农牧户食物消费的影响。研究结果显示:(1)锡林郭勒草地样带生态恢复措施沿样带自北向南实施力度逐渐变弱,草地恢复措施由以草畜平衡、休牧和禁牧为主向休牧和围栏封育为主转变,草地恢复措施中对人类活动的限制也随之由弱变强;(2)沿草地样带自北向南居民食物消费形成了三种模式,在食物消费数量和消费结构上呈显出由牧区特征向农区特征的转变,即人均食物消费量沿样带递减,食物消费由肉蛋奶和蔬菜水果为主向粮食和蔬菜水果为主转变;(3)食物消费模式受到草地供给、收入及其结构、文化习俗和职业分布等生态、经济和社会多方面因素的影响。研究为实现区域生态、经济和社会的协调发展相关政策的制定提供参考。     

Optimal foraging on perilous prey: risk of bill damage reduces optimal prey size in oystercatchers

Intake rate maximization alone is not always sufficient in explainingprey size selection in predators. For example, bivalve-feedingoystercatchers regularly select smaller prey than expected ifthey aimed to maximize their intake rate. It has been proposedthat to these birds large prey are "risky," in the sense thatbirds may damage their bills when feeding on large bivalves.Large bivalves yield more energy, but according to this hypothesisthis is achieved at the expense of energy yield in the longterm when (1) the risk of bill damage increases with prey sizeand (2) foraging with a damaged bill is less effective. In accordancewith this hypothesis, we show that captive oystercatchers feedingon large cockles experienced a high probability of bill tipdamage, while bill damage was absent when cockles were small.Moreover, among free-living oystercatchers the prevalence ofbill damage was correlated with mean cockle size near the capturesite, and the data on captive birds fit in this pattern. Foodintake of captive oystercatchers feeding exclusively on cockleswas reduced by 23% after bill damage, and free-living birdswith damaged bills had 14 g lower mass. Because lower body masswas associated with higher mortality probability, these resultsindicate long-term costs associated with feeding on large cockles.We conclude that the risk of bill damage can potentially explainwhy oystercatchers avoid large bivalves and that oystercatchersmay maximize long-term intake rate by selecting prey sizes thatare "suboptimal" from a short-term rate-maximizing point ofview.     

Towards a synthesis of frameworks in nutritional ecology: interacting effects of protein,carbohydrate and phosphorus on field cricket fitness

Phosphorus has been identified as an important determinant of nutrition-related biological variation. The macronutrients protein (P) and carbohydrates (C), both alone and interactively, are known to affect animal performance. No study, however, has investigated the importance of phosphorus relative to dietary protein or carbohydrates, or the interactive effects of phosphorus with these macronutrients, on fitness-related traits in animals. We used a nutritional geometry framework to address this question in adult field crickets (Gryllus veletis). Our results showed that lifespan, weight gain, acoustic mate signalling and egg production were maximized on diets with different P : C ratios, that phosphorus did not positively affect any of these fitness traits, and that males and females had different optimal macronutrient intake ratios for reproductive performance. When given a choice, crickets selected diets that maximized both lifespan and reproductive performance by preferentially eating diets with low P : C ratios, and females selected diets with a higher P : C ratio than males. Conversely, phosphorus intake was not regulated. Overall, our findings highlight the importance of disentangling the influences of different nutrients, and of quantifying both their individual and interactive effects, on animal fitness traits, so as to gain a more integrative understanding of their nutritional ecology.     

Hydrology‐driven responses of herbivorous geese in relation to changes in food quantity and quality

East Dongting Lake is a Ramsar site and a particularly important wintering ground for herbivorous geese along the East Asian‐Australasian Flyway. The operation of the Three Gorges Dam has changed the water regime and has a significant impact on wetland ecosystems downstream. We studied the responses of two sympatric herbivorous goose species, the Lesser white‐fronted goose Anser erythropus and Bean goose Anser fabalis, to habitat change by investigating their food conditions, habitat selection, and diet composition in the wintering periods of 2016/2017 and 2017/2018, which had early and late water recession, respectively. It was expected that the contrasting water regimes would result in different food conditions and geese responses. The results showed that the food quality and quantity differed significantly between winters. As responses to the high‐quantity/low‐quality food during 2016/2017, more geese switched to feeding on mudflat and exploited plants such as dicotyledons and moss. The tall swards of Carex spp. (dominant plants in the meadow) that developed during the first growing season decreased the food accessibility during the second growing season and hindered the exploitation of newly generated shoots by the geese, which was further confirmed by our clipping control experiment. Nearly all the geese chose to feed on meadow, and Carex spp. made up the majority of their diet in 2017/2018 when there was more low‐quantity/high‐quality food. Compared with the globally vulnerable Lesser white‐fronted geese, the larger‐sized Bean geese seemed to be less susceptible to winter food shortages and exhibited more stable responses. We concluded that the food quality–quantity condition was the external factor influencing the geese responses, while morphological and physiological traits could be the internal factors causing different responses between the two species. This study enhanced the understanding of the influence that habitat change exerts on herbivorous geese in their wintering site in the context of the Three Gorges Dam operation. We suggested that regulating hydrological regime was important in terms of wetland management and species conservation.     

The caloric returns to food collecting: Disruption and change among the Batak of the Philippine tropical forest

The changing economy of a group of tropical forest hunter-gatherers is examined. The Batak of the Philippines today earn many calories by participating in an external market system; once they would have earned these calories within the traditional subsistence system. Gathering and selling Manila copal to purchase rice has partially displaced digging wild yams as the major source of subsistence. Data show that this change has reduced the productive efficiency, of human labor. The Batak must work longer and perform more difficult tasks to obtain the same number of calories by collecting copal as they can obtain by digging wild yams, even as they must today work longer hours anyway to earn necessary cash to satisfy new, nonfood consumer wants. The growing dietary importance of rice is only one aspect of changing food consumption patterns that may be having an unfavorable impact on Batak nutrition.An Arizona State University Faculty Research Grant supported research among the Batak between 1 July and 6 August 1975. A short summary of the research results discussed here appears inBorneo Research Bulletin, April 1976.     

The effect of prey density on foraging mode selection in juvenile lumpfish: balancing food intake with the metabolic cost of foraging

1. In many species, individuals will alter their foraging strategy in response to changes in prey density. However, previous work has shown that prey density has differing effects on the foraging mode decisions of ectotherms as compared with endotherms. This is likely due to differences in metabolic demand; however, the relationship between metabolism and foraging mode choice in ectotherms has not been thoroughly studied. 2. Juvenile lumpfish Cyclopterus lumpus forage using one of two modes: they can actively search for prey while swimming, or they can 'sit-and-wait' for prey while clinging to the substrate using a ventral adhesive disk. The presence of these easily distinguishable foraging modes makes juvenile lumpfish ideal for the study of foraging mode choice in ectotherms. 3. Behavioural observations conducted during laboratory experiments showed that juvenile lumpfish predominantly use the 'cling' foraging mode when prey is abundant, but resort to the more costly 'swim' mode to seek out food when prey is scarce. The metabolic cost of active foraging was also quantified for juvenile lumpfish using swim-tunnel respirometry, and a model was devised to predict the prey density at which lumpfish should switch between the swim and cling foraging modes to maximize energy intake. 4. The results of this model do not agree with previous observations of lumpfish behaviour, and thus it appears that juvenile lumpfish do not try to maximize their net energetic gain. Instead, our data suggest that juvenile lumpfish forage in a manner that reduces activity and conserves space in their limited aerobic scope. This behavioural flexibility is of great benefit to this species, as it allows young individuals to divert energy towards growth as opposed to activity. In a broader context, our results support previous speculation that ectotherms often forage in a manner that maintains a minimum prey encounter rate, but does not necessarily maximize net energy gain.