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OPINION The Value of Animal Behavior in Evaluations of Restoration Success Catherine A. Lindell1,2,3 Abstract Animals are key members of ecosystems, contributing to processes like pollination, seed dispersal, and herbivory. Incorporating measures of animal behavior into evaluations of restoration success will provide critical information that is not available from animal species composition and richness estimates derived from the documentation of species presence and absence. Behavioral studies will (1) allow comparisons of the habitat quality of target and reference sites based on behaviors that have fitness consequences for organisms; (2) provide valuable information about reasons for differences in habitat quality; (3) identify critical resources that make a site suitable or not for Introduction A recent issue of Restoration Ecology has a special section devoted to the measurement of biodiversity (Zerbe & Kreyer 2006) and indices of biodiversity are commonly used to evaluate the success of restoration projects (RuizJaen & Aide 2005). Although faunal components of ecosystems have often been overlooked when assessing the success of restoration efforts (Halle & Fattorini 2004), determining indices of animal biodiversity, usually species composition and richness, is becoming more common (e.g., Jansen 2005; Ottonetti et al. 2006; Summerville et al. 2007). This is a welcome development in that animals are key players in ecosystem function. Arthropods are important herbivores and pollinators (e.g., Coley & Barone 1996; Balvanera et al. 2005) and affect decomposition processes (e.g., Chapman et al. 2003). Birds disperse seeds of numerous plant species (e.g., Wunderle 1997), serve as pollinators and decomposers (Sxekercioğlu 2006) and mediate herbivory levels on plants by foraging on insects (e.g., Greenberg et al. 2000). Mammals disperse seeds (e.g., Bollen et al. 2004) and influence decomposer populations (e.g., Botes et al. 2006). Mollusks and fish have important effects on the abundance and reproductive success of 1 Department of Zoology, 203 Natural Science Building., Michigan State University, East Lansing, MI 48824, U.S.A. 2 Center for Global Change and Earth Observations, 218 Manly Miles Building., Michigan State University, East Lansing, MI 48823, U.S.A. 3 Address correspondence to C. A. Lindell, email [email protected] Ó 2008 Society for Ecological Restoration International doi: 10.1111/j.1526-100X.2008.00389.x JUNE 2008 Restoration Ecology Vol. 16, No. 2, pp. 197–203 particular species; and/or (4) provide information on the mechanisms through which species contribute to ecosystem functions. When resources for the evaluation of restoration success are available, practitioners should carefully consider the costs and benefits of the different variables they could quantify. In some cases, it may be more important to compare the behavior of one or a few critical animal species that contribute to ecosystem function rather than try to document the presence or absence of all species. Key words: animals, behavior, ecosystem function, habitat quality, restoration success. other trophic levels (e.g., Frederiksen et al. 2005; Vaughn et al. 2007). Species composition and richness estimates, however, provide only part of the picture when assessing the responses of animal species to restoration efforts. Behavior is a critical component of the contribution of animals to restoration processes, yet is rarely quantified in evaluations of restoration success. I argue that, in many cases, behavioral sampling in both restoration and reference sites will provide valuable information with which to assess the success of restoration efforts. Documentation of behavior will (1) allow comparisons of the habitat quality of target and reference sites based on behaviors that have fitness consequences for organisms; (2) provide valuable information about reasons for differences in habitat quality; (3) identify critical resources that make a site suitable or not for particular species; and/or (4) provide information on the mechanisms through which species contribute to ecosystem functions. Limitations of Assessing Restoration Success with Presence/Absence Data Successfully restored sites should contribute to individuals being able to reproduce at a rate that allows their population to replace itself, that is, maintain a net reproductive rate equal to or greater than 1. The mere presence of individuals in a site is not strong evidence that the site contributes positively to individual reproductive success and population replacement (Aldridge & Boyce 2007). However, presence/absence data are usually the cornerstones 197 Animal Behavior and Restoration Success of species composition and richness estimates. Previous studies have demonstrated that individuals use habitats in which their reproductive rates are not great enough for population replacement (e.g., Burke & Nol 2000; Hels & Nachman 2002). The most obvious situations in which presence does not reflect the suitability of a habitat are ecological traps. Ecological traps occur when individuals prefer a habitat in which they actually have lower fitness than habitats that are not preferred (Robertson & Hutto 2006). Ecological traps may be particularly common in human-modified landscapes (Battin 2004). Measurements of variables directly related to the likelihood of population replacement will generally be very useful in evaluating the success of restoration efforts. For example, nest success and nest density are strong influences on population replacement in birds (e.g., Pidgeon et al. 2006). However, documenting reproductive success does not explain why individuals in one site have higher reproductive success than individuals in other sites. This missing information may be key to determining components of restoration efforts that are adequate and those that are not. In addition, although some sites undergoing restoration are large enough to provide all the resources necessary for individuals to carry out their daily and seasonal activities, including reproduction, many are not. In these cases, restoration sites may be contributing resources like food or protection from predators but not breeding resources like den or nest sites (e.g., Kus 1998). Thus, even if it were possible to assess individuals’ reproductive success by tracking them to breeding areas outside the restoration sites, it would be difficult to calculate the contribution that the food or protection offered in the restoration site made to the individuals’ reproductive success and to population replacement. Finally, studies to document reproductive success are extremely labor intensive and may yield small sample sizes, particularly for species of special concern (Pejchar et al. 2005). Incorporating studies of behavior into assessments of restoration success will help address these issues. Successful Restoration: A Behavioral Perspective Behavioral patterns often reflect habitat quality (e.g., Vaughan et al. 1996; Johnson 2000; Persson & Stenberg 2006). In addition, an individual’s fitness, that is, the genes it contributes to the next generation (Alcock 2001), is often closely linked to rates and/or outcomes of behaviors (e.g., Good et al. 2000; Persons et al. 2002; Vos & Hemerik 2003; Watson et al. 2007). Recent work indicates that even shortterm behavioral decisions, such as when to leave a food patch, reflect the quality of an individual’s habitat and are linked to potential fitness. For example, fish in high-quality habitats departed from patches of food sooner (i.e., had a higher ‘‘giving up density’’) than fish in low-quality habitats. This variation in behavior explained more than 50% of the variation in fish size, which is positively correlated with potential reproductive output (Persson & Stenberg 2006). Thus, there are strong links between behavior, habi- 198 tat quality, and potential fitness (Olsson et al. 2002). Restoration ecologists can take advantage of these links by quantifying behavior to assess the relative quality of restoration sites and to identify reasons for differences in quality. Quantifying behavior also provides information on potential drivers of species presence or absence, such as the availability or lack of important resources. Finally, documenting behavior can identify how species contribute to ecosystem function and will be particularly useful in cases where determining reproductive success is not feasible because of logistical or financial constraints and for sites that are not used extensively for reproduction. Comparing Habitat Quality Variations in foraging behavior can indicate differences in food availability with higher food availability generally indicating higher quality habitat. Numerous studies with birds and fish have documented higher foraging rates at sites where food is in greater abundance compared with sites with lower food abundance (e.g., Repasky 1996; Delestrade 1999; Shepherd & Boates 1999; Marchand et al. 2002; Wellenreuther & Connell 2002; Kilgo 2005). Observations of foraging-related behaviors can also provide useful information about how differences in habitat quality vary for different species. Two insectivorous bird species in Panama foraged in coffee fields and forest but only one showed a lower foraging rate in coffee, indicating the coffee was lower quality habitat for only this species (Pomara et al. 2003). Anti-predation behavior is also closely related to habitat quality. Pronghorn antelope exhibited greater vigilance rates in habitat near roads, where cars were a common cause of mortality, than further from roads (Gavin & Komers 2006). Increases in predation risk and associated anti-predation behavior can lead to decreases in foraging efficiency (e.g., Metcalfe et al. 1999; Kotler et al. 2004; Orpwood et al. 2006). For example, at greater distances from safe habitat, wild goats spent more time being vigilant and showed higher giving up densities than goats closer to escape habitat (Hochman & Kotler 2007). Thus, in cases where anti-predation behavior and foraging behavior are shown to be strongly correlated, it may be possible to focus on just one of these behaviors in efforts to evaluate the relative quality of reference and restoration sites. To use behaviors such as those described above and in Table 1 as indices of restoration success, it is important to develop a priori predictions about the relationship between the behavior and habitat quality, based on previous empirical work and theory. For example, the relationship between foraging rate and habitat quality may vary between taxonomic groups. Whereas high foraging rates for birds usually indicate high-quality habitat (e.g., Delestrade 1999; Shepherd & Boates 1999; Kilgo 2005), mammalian browsers and grazers may increase food intake rates to compensate for poor quality of available food during some seasons (e.g., Taillon et al. 2006). As another example, recent theory predicts that individuals of solitary species should have Restoration Ecology JUNE 2008 JUNE 2008 Restoration Ecology Spool-and-line Telemetry Birds, mamm Amph, arth, birds, mamm, fish, rept Amph, arth, birds, fish, mamm, moll, rept Amph, mamm Amph, birds, fish, mamm, moll, rept Amph, birds, fish, mamm, moll, rept Amph, arth, birds, fish, mamm, moll, rept Amph, birds, fish, mamm Low Low to Moderate Observations Field experiments High Telemetry Low Low to moderate Mark-recapture Observations High Low Low Moderate Low Low to moderate Telemetry Observations Field experiments Tuberville et al. (2005), Garla et al. (2006) Brashares & Arcese (2002), Walters et al. (2002) Ricketts (2004), Fink et al. (in press) Holl (1998), Zahawi & Augspurger (2006) Lemckert & Brassil (2000) Winker et al. (1995), Baldwin et al. (2006) Grimm & Paill (2001), Wauters et al. (2005) Able et al. (2005), Ortega et al. (2006) Matthysen et al. (1995), Rouquette & Thompson (2007) Wellenreuther & Connell (2002), Pomara et al. (2003) Gavin & Komers (2006) Kotler et al. (2004), Hochman & Kotler (2007) Slabbekoorn & Peet (2003), Sun & Narins (2005) Luck (2003), Stauss et al. (2005) Marsh & Borrell (2001), Yang (2006) References a Groups are those I considered most likely to be of interest in restoration studies and include amphibians (amph), arthropods (arth), birds, fish, mammals (mamm), mollusks (moll), and reptiles (rept). Groups listed are those for which the behavior is regularly measured. b Financial costs associated with equipment, not labor. Social behavior (group size, tendency to group) Visitation rate (in the context of pollination or seed dispersal) Amph, arth, birds, fish, mamm, rept Arth, birds Arth, birds Low to moderate High Observations Site searches, observations Mark-recapture Amph, arth, birds, mamm Vocalizations (frequency, rate) Provisioning behavior Den/nest/oviposition site selection Movement, dispersal (distance, likelihood of) Territory/home range size and overlap Site fidelity (turnover rates) Low to moderate Recordings Birds, fish, mamm Birds, mamm Low Anti-predation (vigilance or allocation of time) Observations Arth, birds, fish, mamm Relative Costb Foraging rate Methods Groupsa Behavior Table 1. Behaviors that could be quantified in studies to evaluate restoration success. References provide information on methods, but were not necessarily conducted in restoration contexts. Animal Behavior and Restoration Success 199 Animal Behavior and Restoration Success larger home range sizes in areas with low habitat quality compared with areas of high habitat quality (McLoughlin et al. 2000; Wauters et al. 2005). However, these patterns may vary with factors such as the sex of the individuals (Lurz et al. 2000). Thus, it is important to consider the species and the context in the design and interpretation of behavioral studies conducted to assess restoration success. Determining Reasons for Differences in Habitat Quality Behavioral observations can provide information that is critical to determining why sites differ in quality. Blue tits in Germany, for example, fledged fewer offspring per nest in poor-quality habitat, a coniferous woodland, compared with high-quality habitat, a deciduous woodland (Stauss et al. 2005). Documentation of this difference, however, provided no information on the reasons for the difference. By also observing parental provisioning behavior, including the distances parents traveled to find food, investigators were able to determine that parents in the poorquality habitat had to travel further per provisioning trip. The number of provisioning trips parents could make in the poor-quality habitat reached an asymptote and limited the number of fledglings produced per brood below that of the high-quality habitat (Stauss et al. 2005). This type of information can inform restoration efforts by identifying how habitat composition and structure influence behaviors that are critical to survival and reproductive success. Identification of Important Resources Behavioral studies demonstrate which resources are important to species. For example, studies of giant barred river frogs in New South Wales revealed that individuals stayed within 20 m of streams and used two types of daytime shelter locations with distinctive characteristics (Lemckert & Brassil 2000). In Hawaii, an endangered Hawaiian honeycreeper foraged primarily on one species of native hardwood tree that can be grown in plantations (Pejchar et al. 2005). The absence or presence of critical resources in sites undergoing restoration may determine the success of the restoration effort with regard to particular species. They may also indicate flaws in current restoration efforts. For example, despite the importance of black locust and hackberry trees to birds in Illinois forests, as demonstrated by behavioral observations, current techniques for savanna restoration in this area discourage the growth of these tree species (Hartung & Brawn 2005). Contributions to Ecosystem Function The Society for Ecological Restoration states that an attribute of a restored ecosystem is that it ‘‘functions normally’’ (The Society for Ecological Restoration 2004). Behaviors that provide functions critical to an ecosystem’s recovery may vary with habitat and/or landscape characteristics that can be considered in the design of restoration efforts. Several studies have demonstrated that bird behavior is influenced by restoration design, including the size of planted 200 patches (Zahawi & Augspurger 2006; Fink et al. in press), the types of perches available (Holl 1998), and the tree species planted (Zahawi & Augspurger 2006; Fink et al. in press). Variations in behavior are linked to variation in processes like seed dispersal that contribute to restoration success (Holl 1998; Zahawi & Augspurger 2006). Animal behavior influences other processes that are critical to restoration success and ecosystem function including decomposition, herbivory, and pollination (Chapman et al. 2003; Van Bael & Brawn 2005; Feldman 2006). The design of restoration projects may facilitate or impede the recovery of these processes through effects on animal behavior. Apportioning Resources to Behavioral Studies Resources to evaluate the success of restoration efforts are often very limited. Thus, when resources for evaluation are available, practitioners should carefully choose variables that are likely to provide them with the greatest amount of information. Although determining species presence/absence and species richness are commonly measured and important in many cases, particular species often play more significant ecological roles than others and/or are of greater conservation concern than others. For example, one or a few species may be particularly important seed dispersers or pollinators in a system (e.g., Wenny & Levey 1998; Ricketts 2004). Thus, a more strategic use of resources in some cases may be to measure the behavior of a few key species, rather than to try to comprehensively sample the presence of all species. Methods used to quantify behavior vary in equipment costs. Observations of foraging or vigilance behavior, for example, could be incorporated into a study for a low cost—in many cases, a pair of binoculars. At the other end of the spectrum, the equipment necessary to document movement patterns with telemetry is very expensive. Table 1 lists some of the behaviors that could be measured in studies to evaluate restoration success and their relative costs with regard to equipment. Another cost of incorporating behavioral assessments into restoration studies is finding or developing the expertise to conduct such assessments. Partnering with behavioral ecologists is one way for restoration ecologists to gain access to this expertise (e.g., Fink et al. in press). Another way is to take advantage of local amateur knowledge. Members of birding and nature clubs are often eager to share their know-how about animal behavior with newcomers and, with some effort, it is not difficult to develop observation skills that are sufficient to identify a few key species and categorize their behaviors. A final cost to behavioral assessments is the labor involved in conducting the studies. Quantifying most of the behaviors in Table 1 would involve moderate to high labor costs. A potential source of labor is volunteers. Volunteers have been used successfully to gather data in animal conservation and/or monitoring programs in a number of countries (e.g., Garnett et al. 1999; Gregory et al. 2005; Restoration Ecology JUNE 2008 Animal Behavior and Restoration Success de Solla et al. 2006) and the Cornell Lab of Ornithology currently has several programs that use volunteers to collect data on behavior (Cornell Lab of Ornithology 2008). Given the strong interest many volunteers have in habitat conservation (Weston et al. 2003), studies involving monitoring in restoration sites would be attractive. Administering volunteer programs is not effortless, but can produce large quantities of useful data and involve local citizens in science (West 2000). Volunteer labor will not work in all situations, for example, in areas of the developing world where people struggle to make a living. Conclusions Few previous studies have incorporated behavioral data into assessments of restoration success (e.g., Holl 1998; Brusati et al. 2001; Armitage et al. 2007). I argue that measurements of behavior should regularly be considered by investigators as components of evaluations of success. Quantification of behavior often will provide a better index of the quality of a site than the simple presence or absence of species and will provide insight as to why sites differ in quality. Behavioral studies also will indicate resources that are important to species. They will provide information as to how species contribute to ecosystem function and how these contributions vary with restoration design. Behavior will often demonstrate the mechanisms through which the attributes of restored ecosystems, including a characteristic assemblage of species and a normally functioning ecosystem (The Society for Ecological Restoration 2004), are realized. Given the importance of restoration in a world with vast areas of degraded lands (Lamb et al. 2005), it makes sense to make the most of our limited resources by choosing the most information-rich ways to evaluate restoration success. Implications for Practice Animals are key players in many ecosystem processes yet often receive little attention in evaluations of restoration success. d When the animal components of restoration efforts are assessed, it is often with species composition and richness estimates derived from presence/absence data. d Measures of animal behavior can compensate for many of the limitations of presence/absence data by providing information about why reference and restoration sites differ in quality, identifying critical resources for animals, and documenting how animals contribute to ecosystem function. d When resources are available to assess the success of restoration efforts, behavioral studies should be considered as a potentially rich source of information. d If practitioners choose to use animal behavior to evaluate restorations success, they should carefully develop testable predictions to guide the design, analysis, and interpretation of such studies. d JUNE 2008 Restoration Ecology Acknowledgments I am grateful to Karen Holl, Richard Hobbs, Emily Morrison, and Rakan Zahawi for helpful comments on earlier drafts of this manuscript. 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