Primate foraging

Primates often use mental maps to remember the location of food resources, which allows them to plan goal-oriented paths to these resources. {{Cite journal |last1=Erhart |first1=Elizabeth M. |last2=Overdorff |first2=Deborah J. |date=2008-01-07 |title=Spatial Memory during Foraging in Prosimian Primates: Propithecus edwardsi and Eulemur fulvus rufus |url=https://karger.com/fpr/article-abstract/79/4/185/143445/Spatial-Memory-during-Foraging-in-Prosimian |journal=Folia Primatologica |volume=79 |issue=4 |pages=185–196 |doi=10.1159/000112911 |pmid=18176079 |issn=0015-5713|url-access=subscription }} This spatial memory is crucial in environments where food availability is seasonal and unpredictable. {{Cite journal |last1=Trapanese |first1=Cinzia |last2=Meunier |first2=Hélène |last3=Masi |first3=Shelly |date=2019 |title=What, where and when: spatial foraging decisions in primates |url=https://onlinelibrary.wiley.com/doi/10.1111/brv.12462 |journal=Biological Reviews |language=en |volume=94 |issue=2 |pages=483–502 |doi=10.1111/brv.12462 |pmid=30211971 |issn=1469-185X|url-access=subscription }}{{Cite journal |last=Stone |first=Anita I. |date=2007 |title=Responses of squirrel monkeys to seasonal changes in food availability in an eastern Amazonian forest |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajp.20335 |journal=American Journal of Primatology |language=en |volume=69 |issue=2 |pages=142–157 |doi=10.1002/ajp.20335 |pmid=17154390 |issn=1098-2345|url-access=subscription }} The ability to integrate spatial and temporal information is less understood, but it is suggested that primates may learn temporal patterns of food availability to enhance foraging efficiency.

{{Main|Optimal foraging theory}}

Primate foraging strategies are heavily influenced by the need to balance nutrient intake. Theories such as energy maximization and minimization have been proposed to explain their dietary choices. {{Cite journal |last=Righini |first=Nicoletta |date=2017 |title=Recent advances in primate nutritional ecology |url=https://onlinelibrary.wiley.com/doi/10.1002/ajp.22634 |journal=American Journal of Primatology |language=en |volume=79 |issue=4 |pages=e22634 |doi=10.1002/ajp.22634 |pmid=28076653 |issn=1098-2345|url-access=subscription }}

Primates living in seasonal or fragmented environments must balance energy expenditure with food intake. This typically results in different foraging strategies, commonly categorized as either energy maximization, expanding space use to seek high-quality foods, or energy minimization, reducing movement and activity to conserve energy during periods of scarcity. {{Cite journal |last=Schoener |first=Thomas W. |date=1971 |title=Theory of Feeding Strategies |url=https://www.jstor.org/stable/2096934 |journal=Annual Review of Ecology and Systematics |volume=2 |issue=1 |pages=369–404 |doi=10.1146/annurev.es.02.110171.002101 |jstor=2096934 |bibcode=1971AnRES...2..369S |issn=0066-4162|url-access=subscription }}

Some examples of those strategies being adopted can be seen in Black-fronted titi monkeys (Callicebus nigrifrons) that they minimize their energy use during low fruit availability by decreasing travel distances and diversifying their diet. {{Cite journal |last1=Nagy-Reis |first1=Mariana B. |last2=Setz |first2=Eleonore Z. F. |date=2017-01-01 |title=Foraging strategies of black-fronted titi monkeys (Callicebus nigrifrons) in relation to food availability in a seasonal tropical forest |url=https://link.springer.com/article/10.1007/s10329-016-0556-9 |journal=Primates |language=en |volume=58 |issue=1 |pages=149–158 |doi=10.1007/s10329-016-0556-9 |pmid=27485746 |issn=1610-7365|url-access=subscription }} In a fragmented landscape, Alouatta caraya groups adopted different strategies: island groups relied on fallback foods and minimized space use, while mainland groups expanded their ranges.{{Cite journal |last=Ludwig |first=Gabriela |date=2006 |title=Área de vida e uso do espaço por Alouatta caraya (Humboldt, 1812) em ilha e continente do Alto Rio Paraná / |url=https://acervodigital.ufpr.br/handle/1884/4717 |journal=OAI:ufpr.br:221524 |language=pt}} Propithecus diadema, a folivorous lemur, increases movement during low food availability, while Eulemur fulvus, a frugivore, expand its range to maximize food intake.{{Cite journal |last1=Rahalinarivo |first1=Vololonirina |last2=Rakotomanana |first2=Hajanirina Fanomezantsoa |last3=Randrianasy |first3=Jeannot |last4=Ranaivoarisoa |first4=Jean Freddy |last5=Ramorasata |first5=Bruno |last6=Raharison |first6=Jean Luc Fanomezantsoa |last7=Irwin |first7=Mitchell |date=2023 |title=Activity budget and seasonal activity shifts in sympatric lemurs: Increased lean season effort in a cathemeral frugivore contrasts with energy conservation in a diurnal folivore |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajp.23556 |journal=American Journal of Primatology |language=en |volume=85 |issue=12 |pages=e23556 |doi=10.1002/ajp.23556 |pmid=37779335 |issn=1098-2345}} Cebus olivaceus in Venezuela expand their home range during food shortages, pursuing an energy maximization strategy.{{Cite journal |last=Robinson |first=J. G. |date=1988-09-01 |title=Group size in wedge-capped capuchin monkeys Cebus olivaceus and the reproductive success of males and females |url=https://link.springer.com/article/10.1007/BF00300353 |journal=Behavioral Ecology and Sociobiology |language=en |volume=23 |issue=3 |pages=187–197 |doi=10.1007/BF00300353 |bibcode=1988BEcoS..23..187R |issn=1432-0762|url-access=subscription }} Cebus capucinus in Costa Rica expand their space use during scarcity, but reduce movement near water sources during hotter months {{Cite journal |last1=Campos |first1=Fernando A. |last2=Fedigan |first2=Linda M. |date=2014-05-01 |title=Spatial ecology of perceived predation risk and vigilance behavior in white-faced capuchins |url=https://academic.oup.com/beheco/article-abstract/25/3/477/510312 |journal=Behavioral Ecology |volume=25 |issue=3 |pages=477–486 |doi=10.1093/beheco/aru005 |issn=1045-2249}}{{Citation |last1=Gómez-Posada |first1=Carolina |title=Ranging Responses to Fruit and Arthropod Availability by a Tufted Capuchin Group (Sapajus apella) in the Colombian Amazon |date=2019 |work=Movement Ecology of Neotropical Forest Mammals: Focus on Social Animals |pages=195–215 |editor-last=Reyna-Hurtado |editor-first=Rafael |url=https://link.springer.com/chapter/10.1007/978-3-030-03463-4_12 |access-date=2025-04-03 |place=Cham |publisher=Springer International Publishing |language=en |doi=10.1007/978-3-030-03463-4_12 |isbn=978-3-030-03463-4 |last2=Rey-Goyeneche |first2=Jennifer |last3=Tenorio |first3=Elkin A. |editor2-last=Chapman |editor2-first=Colin A.|url-access=subscription }}

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Social factors, such as feeding competition and dominance hierarchies, play a significant role in primate foraging behavior.{{Cite journal |last1=Foerster |first1=Steffen |last2=Cords |first2=Marina |last3=Monfort |first3=Steven L. |date=2011 |title=Social behavior, foraging strategies, and fecal glucocorticoids in female blue monkeys (Cercopithecus mitis): potential fitness benefits of high rank in a forest guenon |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajp.20955 |journal=American Journal of Primatology |language=en |volume=73 |issue=9 |pages=870–882 |doi=10.1002/ajp.20955 |pmid=21495049 |issn=1098-2345|url-access=subscription }} For instance, in some species, females may have priority access to food resources, which can influence group dynamics and social structures. {{Cite journal |last=Box |first=Hilary O. |date=1997-02-14 |title=Foraging Strategies among Male and Female Marmosets and Tamarins (Callitrichidae): New Perspectives in an Underexplored Area |url=https://brill.com/view/journals/ijfp/68/3-5/article-p296_15.xml |journal=Folia Primatologica |volume=68 |issue=3–5 |pages=296–306 |doi=10.1159/000157255 |pmid=9360311 |issn=1421-9980|url-access=subscription }} The Marginal Value Theorem has been applied to understand how primates make decisions about when to leave a food patch, balancing the rate of resource intake with the time spent in a patch. {{Cite journal |last1=Turrin |first1=Courtney |last2=Fagan |first2=Nicholas A. |last3=Dal Monte |first3=Olga |last4=Chang |first4=Steve W. C. |date=2017-09-12 |title=Social resource foraging is guided by the principles of the Marginal Value Theorem |journal=Scientific Reports |language=en |volume=7 |issue=1 |pages=11274 |doi=10.1038/s41598-017-11763-3 |issn=2045-2322 |pmc=5596022 |pmid=28900299|bibcode=2017NatSR...711274T }}

In many primate species, juveniles develop foraging skills by observing and mimicking the behavior of older or more experienced group members. This social learning process helps young individuals identify safe and nutritious foods, increasing their foraging efficiency over time. {{Cite journal |last1=Rapaport |first1=Lisa G. |last2=Brown |first2=Gillian R. |date=2008 |title=Social influences on foraging behavior in young nonhuman primates: Learning what, where, and how to eat |url=https://onlinelibrary.wiley.com/doi/10.1002/evan.20180 |journal=Evolutionary Anthropology: Issues, News, and Reviews |language=en |volume=17 |issue=4 |pages=189–201 |doi=10.1002/evan.20180 |issn=1520-6505|url-access=subscription }}

Group size in primates is often limited by the balance between the time available for foraging and the energetic benefits of food patches. When groups become too large, competition can increase travel costs and reduce foraging efficiency, leading to fission–fusion behavior as a strategy to manage these constraints. {{Cite journal |last=Grove |first=Matt |date=2012-02-01 |title=Space, time, and group size: a model of constraints on primate social foraging |url=https://www.sciencedirect.com/science/article/abs/pii/S0003347211005033 |journal=Animal Behaviour |volume=83 |issue=2 |pages=411–419 |doi=10.1016/j.anbehav.2011.11.011 |issn=0003-3472|url-access=subscription }}

File:Stone tool use by a capuchin monkey.jpg

Extractive foraging, which involves locating and processing embedded foods, is a strategy used by some primates, such as capuchin monkeys{{Cite journal |last1=Ottoni |first1=Eduardo B. |last2=Izar |first2=Patrícia |date=2008 |title=Capuchin monkey tool use: Overview and implications |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/evan.20185 |journal=Evolutionary Anthropology: Issues, News, and Reviews |language=en |volume=17 |issue=4 |pages=171–178 |doi=10.1002/evan.20185 |issn=1520-6505|url-access=subscription }} and chimpanzees.{{Cite journal |last=Watts |first=David P. |date=2008 |title=Tool Use by Chimpanzees at Ngogo, Kibale National Park, Uganda |url=https://link.springer.com/article/10.1007/s10764-007-9227-4 |journal=International Journal of Primatology |language=en |volume=29 |issue=1 |pages=83–94 |doi=10.1007/s10764-007-9227-4 |issn=1573-8604|url-access=subscription }} This behavior is linked to cognitive abilities and may involve the use of tools. {{Cite journal |last=King |first=B. J. |date=1986 |title=Extractive foraging and the evolution of primate intelligence |url=https://link.springer.com/article/10.1007/BF02436709 |journal=Human Evolution |language=en |volume=1 |issue=4 |pages=361–372 |doi=10.1007/BF02436709 |issn=1824-310X|url-access=subscription }} While extractive foraging is not unique to primates, it is suggested that the complexity of these behaviors in primates may have contributed to the evolution of their intelligence.

Chimpanzees can use tools to dig up underground food, showing behaviors once thought unique to humans. In experiments, they spontaneously selected and reused tools to excavate buried items. {{Cite journal |last1=Motes-Rodrigo |first1=Alba |last2=Majlesi |first2=Parandis |last3=Pickering |first3=Travis Rayne |last4=Laska |first4=Matthias |last5=Axelsen |first5=Helene |last6=Minchin |first6=Tanya C. |last7=Tennie |first7=Claudio |last8=Hernandez-Aguilar |first8=R. Adriana |date=2019-05-15 |title=Chimpanzee extractive foraging with excavating tools: Experimental modeling of the origins of human technology |journal=PLOS ONE |language=en |volume=14 |issue=5 |pages=e0215644 |doi=10.1371/journal.pone.0215644 |doi-access=free |issn=1932-6203 |pmc=6519788 |pmid=31091268|bibcode=2019PLoSO..1415644M }} But also, tool-assisted foraging behaviors in chimpanzees, such as ant-dipping or termite-fishing, often depend on the availability of resources and encounter rates, not just cognitive capacity.{{Cite journal |last1=Koops |first1=Kathelijne |last2=McGrew |first2=William C. |last3=Matsuzawa |first3=Tetsuro |date=2013-01-01 |title=Ecology of culture: do environmental factors influence foraging tool use in wild chimpanzees, Pan troglodytes verus? |url=https://www.sciencedirect.com/science/article/abs/pii/S0003347212004824 |journal=Animal Behaviour |volume=85 |issue=1 |pages=175–185 |doi=10.1016/j.anbehav.2012.10.022 |issn=0003-3472|url-access=subscription }}

The interaction between ecological variables and morphological traits influences primate foraging adaptations. For example, the physical ability to process certain types of food can dictate dietary choices and foraging strategies. {{Cite journal |last1=Janson |first1=Charles H. |last2=Boinski |first2=Sue |date=1992 |title=Morphological and behavioral adaptations for foraging in generalist primates: The case of the cebines |url=https://onlinelibrary.wiley.com/doi/abs/10.1002/ajpa.1330880405 |journal=American Journal of Physical Anthropology |language=en |volume=88 |issue=4 |pages=483–498 |doi=10.1002/ajpa.1330880405 |pmid=1503120 |issn=1096-8644|url-access=subscription }} Mandible shape in primates is linked to how they chew different foods. {{Cite journal |last1=Ross |first1=Callum F. |last2=Iriarte-Diaz |first2=Jose |last3=Nunn |first3=Charles L. |date=2012-06-01 |title=Innovative Approaches to the Relationship Between Diet and Mandibular Morphology in Primates |url=https://link.springer.com/article/10.1007/s10764-012-9599-y |journal=International Journal of Primatology |language=en |volume=33 |issue=3 |pages=632–660 |doi=10.1007/s10764-012-9599-y |issn=1573-8604|url-access=subscription }}

Primate body size influences foraging strategies. Small primates prioritize high-energy, easy-to-digest foods such as insects, nectar, and ripe fruits, while larger primates can process more fibrous, lower-quality foods like mature leaves.{{Cite journal |last=Garber |first=P. A. |date=1987 |title=Foraging Strategies among Living Primates |url=https://www.jstor.org/stable/2155875 |journal=Annual Review of Anthropology |volume=16 |pages=339–364 |doi=10.1146/annurev.an.16.100187.002011 |jstor=2155875 |issn=0084-6570|url-access=subscription }} In high-altitude Himalayan habitats, gray langurs shift from preferred foods (young leaves, fruits) to fibrous, less-profitable fallback foods (bark, mature leaves, roots) during the winter, showing seasonal dietary flexibility despite morphological constraints. {{Cite journal |last1=Sayers |first1=Ken |last2=Norconk |first2=Marilyn A. |date=2008-04-01 |title=Himalayan Semnopithecus entellus at Langtang National Park, Nepal: Diet, Activity Patterns, and Resources |url=https://link.springer.com/article/10.1007/s10764-008-9245-x |journal=International Journal of Primatology |language=en |volume=29 |issue=2 |pages=509–530 |doi=10.1007/s10764-008-9245-x |issn=1573-8604|url-access=subscription }}

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