A Practical Technique for Measuring the Behavior of Foraging Animals
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g k sTHow A PractIcal Technique for )D Do Measuring the Behavior of It _ . It Foraging hnlmalS RosemaryJ. Smith Joel S. Brown Animal behavior is well suited for havioral questions that can be readily ness (survivaland reproduction).If so, illustrating ecological and evolution- addressed gives students the opportu- an animal should leave a patch when ary principles. For ecology projects or nity to suggest their own hypotheses its benefits of continued feeding in the labs, animal behavior has a number of and experiments.The projectsempha- patch no longer exceed its cost of feed- advantages:It is inherently interesting size the methodology of a scientist ing in the patch. The formal frame- Downloaded from http://online.ucpress.edu/abt/article-pdf/53/4/236/45243/4449276.pdf by guest on 01 October 2021 to most students, it can be observed, it and, we hope, the excitement that work behind the assertion in the pre- is flexible and it can change quickly. accompanies creative thinking and vious sentence is called Optimal Unfortunately, meaningful observa- novel results. Foraging Theory (see Stephens & tions of behavior tend to be time- Krebs 1986). consuming and difficultto analyze. To We begin by elaborating a rule for overcome this handicap, we introduce Rationale when an animal should leave a patch an indirect procedure that uses the and then discuss how this rule can be foraging behavior of animals at exper- "Talking" to animals requires a common language, and feeding be- used to reveal aspects of that animal's imental food patches to address ques- ecology. An example from our own tions in animal behavior. The data are havior can provide just such a lan- guage. Mixing food into a substrate work illustratesthis rule. We provided the food left behind by the animals. kangaroo rats living in the Sonoran Such creates a food patch and provides a data are quantitative and easily Desert of Arizona with shallow alumi- analyzed using a variety of technique for "asking" foragers how descriptive num pans (45 x 45 x 2.5 cm) filled with or inferential statistics. The experi- they perceive their environment. The a known amount of millet seed (3 g) mental procedure encourages stu- food patches are like a survey distrib- dents to ask questions relevant to how uted to the animalsbeing investigated. that has been mixed into 3 1 of sifted environmentalfactors such as climate, The animals fill out the survey by dirt. Kangaroorats foraged enthusias- vegetation structure and predation digging for and harvesting food from tically in these food patches. As more risk influence animals'foraging behav- the tray's substrate and then sign and more seeds are eaten, the large ior. When several species are included these trays with footprints, body amount of dirt in the patch makes it within the study, interspecific differ- marks and other telltale disturbances increasingly difficult for the kangaroo ences in behavior provide an easy and of the substrate. The amount of food rats to find the seeds that remain. relevant introduction to questions of left in the patch contains the answer to Therefore, the benefit of remaining in niche separation and species coexis- the survey. the patch declines. At some density of tence. The following approach to feeding seeds the cost of looking for the next Because it uses food left by animals behavior represents evolutionaryecol- seed exceeds the benefit derived from in feeding patches, the technique does ogy. From an ecological perspective, harvesting and consuming that seed. not require hours of observation, han- two universal properties of life in- At this criticalseed density the kanga- dling of animals or expensive equip- clude: roo rat should leave the patch. If it ment. Because feeding is central to all 1. Births and deaths continues to forage past this density, it animals regardless of environment, 2. The need for energy and nutri- forages at a loss (costs exceed bene- studies can be conducted on school ents to maintainhomeostasis and fits). If it quits early, it forfeits the food campuses using urban birds (spar- life functions that it could have profitablygained. rows, doves, pigeons, starlings) or An animal should leave a food patch small mammals (squirrels, mice and From an evolutionary perspective, when its benefits equal its costs (see rats). The cost/benefit theory behind these two properties of life encourage Brown 1988 for a formal treatment of the technique is intuitive and accessi- animals to have feeding behaviorsthat the theory). To apply this foraging ble to students. The breadth of be- allow them to acquire food in an effi- theory, the animal must experience a cient manner. declining rate of return from spending Feeding animals make decisions time in the food patch, and the animal about where to feed (habitatselection, must experience both benefits and Rosemary J. Smith teaches in the de- MacArthur & Pianka 1966), what to costs from exploiting the patch. The partment of ecology and evolutionary feed on (diet choice, Pulliam1974) and benefit to biology at the Universityof Arizona,Tuc- the forager is its rate of son, AZ85721. Joel S. Brownteaches in when to quit an area and move to harvest of food times (H) considered the department of biological sciences another location (patch use, Charnov in terms of energy or nutrient intake. at the Universityof Illinois,Box 4348, 1976). We assume that natural selec- The costs of spending time in a food Chicago, IL60680. tion has favored those animals whose patch include energetic costs (E) of feeding decisions maximize their fit- searchingfor and handling food items, 236 THEAMERICAN BIOLOGY TEACHER, VOLUME 53, NO. 4, APRIL1991 walk. And finally, if you knew there VARYINGFORAGING COSTS was another full jar on the shelf, or [ENERGETIC,MISSED OPPORTUNITY OR PREDATIONI that you were missing out on a great TV show (high missed opportunity costs), again you would leave more peanut butter in the jar. m The amount of peanut butter left in the jar, the GUD, is a clearreflection of D z 4- your perceptionof these three types of costs. Refer back to the equation and note that if any two costs are held constant, any differencesin GUDs be- o 3 tween experimentaltreatments will in- dicate the effect of the variationin the third cost (see Figure 1 for summary). 2 z Methods Using GUDs to Study Feeding Downloaded from http://online.ucpress.edu/abt/article-pdf/53/4/236/45243/4449276.pdf by guest on 01 October 2021 o 1 Costs & Opportunities The model predicts that two identi- > Low HIGH cal feeding patches placed side by side within the same habitat should have Figure 1. An illustration of how giving up density of peanut butter might vary the same GUD. This is because both with experimental conditions. The LOW giving up density relative to HIGH trays have the same substrate and indicates a lower energetic, missed opportunity or predation cost of extracting food items (controls for H) are subject peanut butter from a jar. to the same microclimaticconditions (controlsfor E), are equally exposed to potential dangers (controls for P), are equally situated for alternative activi- ties (controls for M) and are available A Useful Analogy of the Rule for to the same animals (differentanimals risk of predation (P) and the missed in their of the Leaving Patches may vary perceptions opportunity cost (M) of not engaging food, energetic costs of foraging, pre- in alternativefitness-enhancing activi- We have found that the following dation risk and opportunities pro- ties such as mating, defending a terri- human example of the patch-leaving vided by alternative activities). The tory or-exploiting a richer food patch. rule is useful because it provides stu- above prediction allows for controlled The patch-leaving rule can then be dents with a mental image of the way manipulations of the model's compo- written as: GUDs change when the cost compo- nents (H, E, P and M). The effect of nents are manipulated. Imagine an any one of the components on the H = E + P + M almost empty peanut butterjar (this is GUD can be tested by holding the the resource patch). As you remove other components constant. In what This rule provides the model for peanut butter from the jar it becomes follows it is important to remember investigating foraging behavior. An more and more difficultto get the next that higher GUDs mean more food is spoonful out (this demonstrates a de- left in the patch, hence the animals animal should leave a food patch clining rate of return). According to either perceived higher costs or lower when the harvest rate from the patch the rule, you will quit removing pea- benefits from harvesting the patch. no longer exceeds the sum of the en- nut butter when the benefit (the har- Consider an investigation of factors ergetic, predation and missed oppor- vest rate) just equals the sum of the that may influence harvest rates. Plac- tunity costs of foraging. In reality, it is costs (energetic,predation and missed ing food patches side by side within difficult to measure H and even more opportunity). The amount of peanut the same habitat can control for ener- difficult to measure H, E, P and M in butter left in the jar will be your GUD. getic costs, predation risk and missed terms of their fitness consequences, Now, what happens when one of opportunity costs. By varying food which is their true common currency. the costs is varied while the others are types, food size or substrate, it is pos- Fortunately, the density of food left held constant? When it is cold and sible to test for their effects on H. behind in a patch when an animal more energy is required to scrape out Larger amounts of substrate or more quits foraging is a good surrogate for the last bit of peanut butter (higher difficult substrates should increase H.