Vol. 29: 69–79, 2015 ENDANGERED SPECIES RESEARCH Published online November 25 doi: 10.3354/esr00700 Endang Species Res OPENPEN ACCESSCCESS Conservation implications of harpy eagle Harpia harpyja predation patterns Everton B. P. Miranda* Programa de Pós-graduação em Ecologia e Conservação da Biodiversidade, Universidade Federal do Mato Grosso, Av. Fernando Corrêa da Costa, no. 2367 - Bairro Boa Esperança, Cuiabá − MT, CEP 78060-900, Brazil ABSTRACT: Knowledge of the food habits of threatened taxa is key for their effective conserva- tion, especially in top predators where prey species are frequently also hunted by humans. The harpy eagle Harpia harpyja is the largest living eagle, and is considered Near Threatened by the IUCN. Its main threats are persecution by humans and habitat loss. Predation patterns of this spe- cies have been the subject of several descriptive studies, each reflecting the idiosyncrasies of the study area. Systematizing these data permits a transition from descriptive treatments of harpy food habits to a predictive focus, based on defensive prey strategies and foraging theory. This gen- erates information that can enhance management and conservation decisions. Literature data were summarized and standardized, allowing comparison between studies. Results indicate that harpy eagles feed mainly on sloths and other prey with passive antipredator strategies, with sloths accounting for 50% of prey items and biomass consumed. Large monkeys such as howlers (Alouatta spp.) and capuchins (Sapajus and Cebus spp.) are the next most important prey, but combined, primates form only ~20% of the consumed prey biomass. Predation seldom occurs on animals weighing more than 5 kg. This is positive from a conservation point of view, since sloths are not game species, precluding competition between harpy eagles and subsistence hunting. KEY WORDS: Raptor · Bradypus · Choloepus · Alouatta · Prey defenses · Top predator · Diet INTRODUCTION such studies may offer little to improve the effective management of this type of species. Knowledge of the food habits of threatened taxa is As predators with a large body size, raptors have essential for their effective conservation (Harper et low densities and high resource needs, both of which al. 2006, Real et al. 2009). This is especially true for are strong predictors of high extinction risk (Krüger & apex predators, since they are commonly dependent Radford 2008, Lees & Peres 2008). The largest-ever on large-bodied prey, which, in turn, are frequently raptor, the Haast eagle Harpagornis moorei, weighed hunted by humans (Hayward et al. 2012, Lyngdoh et up to 17 kg, preyed on giant moa and disappeared al. 2014). Effective conservation planning (e.g. re in - shortly after Maori colonization of New Zealand tro ductions, thematic conservation units, prevention around 1400, probably due to prey loss, direct perse- of livestock losses) for such species must therefore cution, or both (Holdaway 1991, Scofield & Ashwell include precise assessment of prey base composition. 2009). This kind of extinction event has occurred Predation patterns can be affected by a wide range of repeatedly on islands, and has affected both terres- ecological constraints that vary across the geo- trial and flying giant raptors, such as Ornimegalonyx graphic range of the predator species. Thus, local oteroi in Cuba (Arredondo 1976) and Stephanoaetus food habit descriptions may well have little practical mahery in Madagascar (Goodman 1994). The Philip- utility in a range-wide conservation framework (Hay- pine eagle Pitheco phaga jefferyi (4.4−6 kg; Gamauf ward & Kerley 2005, Schweiger et al. 2015). Other et al. 1998), the second largest raptor on earth, is cat- than local descriptions of prey-base composition, egorized as Critically Endangered by the IUCN; no © The author 2015. Open Access under Creative Commons by *Corresponding author: [email protected] Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited. Publisher: Inter-Research · www.int-res.com 70 Endang Species Res 29: 69–79, 2015 more than 250 pairs remain in the wild, and this spe- harpy eagle diet, in order to answer the following cies is faced with prey depletion, habitat loss, and questions: (1) What are the main prey items of harpy poaching (Birdlife International 2013a). eagles, and what is their proportional biomass contri- The harpy eagle Harpia harpyja is the world’s largest bution to the diet? (2) Data on how many prey must living eagle, weighing between 4.8 and 7.6 kg (Sick be collected in order to adequately represent feeding 1984, Ferguson-Lees & Christie 2001). In Central and habits? Additionally, 2 hypotheses, based on predic- South America, harpy eagles are threatened by retali- tions from foraging theory, were tested: (1) the pro- ation for predation (imagined and real) on domestic portion of sloths in the diet will negatively affect animals, habitat loss, use as food, and by curious set- niche width, and (2) the antipredator strategy of the tlers and colonists who want to see the birds closer at main prey species will be passive, with morphologi- hand (Trinca et al. 2008, Godoi et al. 2012, Freitas et cal and behavioral specializations for crypsis. Such al. 2014). The species is distributed over forest ecosys- information can be used to help predict predation tems in Central and South America, but has nearly patterns and carrying capacity of unstudied popula- vanished from Cerrado and Atlantic Forest environ- tions, and have implications for viable and effective ments (De Oliveira & Silva 2006, Aguiar-Silva et al. management and conservation strategies. 2012, Silva et al. 2013). Harpy eagles are categorized as Near Threatened by the IUCN, with declining pop- ulation trends (Birdlife International 2013b). METHODS Foraging theory predicts that predators with low search costs act to maximize energy gain by preying Data acquisition and compilation on animals of low detectability, but with high catcha- bility once detected (Stephens et al. 2007). In contrast A data search was made with Google Scholar using to most large raptors, harpy eagles rarely soar, and Harpia harpyja and the following keywords: harpy prey searching is dependent on visual and hearing eagle, arpía, gavião-real, harpia combined with diet, skills, with relatively low energy expenditure while food habits, habitos alimentarios, and dieta. This hunting (Touchton et al. 2002). This suggests (sensu allowed me to access published and unpublished foraging theory) that prey such as sloths (Megalony- studies in English, Portuguese, and Spanish. Studies chidae and Bradypodidae), which rely mainly on which were known to exist but which could not be crypsis to avoid predation, are likely to be important downloaded were obtained in direct contact with prey for harpy eagles. Prey that practice flight as well authors or by contacting the Brazilian Harpy Eagle as crypsis can increase manipulation and/or search- Conservation Program (gaviaoreal.inpa.gov.br). ing time (Eason 1989, Touchton et al. 2002, Ferrari & I tabulated relevant data collected by direct Port- Carvalho 2003). In the context of top-down the- observation or from analyses of bone material and ory, and since raptors strongly shape primate behav- pellets collected from beneath nest trees or from ior (Gil-da-Costa et al. 2003, Willems & Hill 2009), the inside nests accessed by climbing. I considered only absence of harpy eagles can indirectly increase her- studies whose estimation of the minimum number of bivore pressure on vegetation (Terborgh et al. 2001, prey individuals included standardized collection Ori huela et al. 2014) via population growth and in - proce dures or studies that counted skulls, pelvises, creased vegetation consumption. and other unpaired bones or perfectly paired limb Although several excellent studies have investigated bones. This procedure permits the use of unpub- prey composition by harpy eagles, all of them were lished studies when data were derived from robust restricted by local idiosyncrasies, such as regio nal prey and standardized methods. Two studies (Chebez et abundances and habitat types. Hence, an overview of al. 1990, Anfuso et al. 2008) were excluded because predation patterns over the entire distribution of harpy the sample sizes of prey items were small (<5) and eagles is timely. Aguiar-Silva et al. (2014) conducted a uncertainties in prey item identification. A further 2 review in order to compare their results with pub- studies (Rettig 1978, Izor 1985) were collapsed into lished literature, but they used political rather than a single sample since they came from a single nest- habitat boundaries and did not distinguish between ing event investigated at different times. When pos- multiple nests in a single sample (e.g. Fowler & Cope sible, I considered each individual bird as a repli- 1964, Muniz-Lopez et al. 2007) or multiple studies of a cate, but most studies used each nest as an single nest (e.g. Rettig 1978, Izor 1985). independent unit, and others collapsed several nests In the present study, I synthetized the available into 1 data set. In those cases, nest data were com- information from a variety of previous studies on puted as a single replicate. Miranda: Harpy eagle predation patterns 71 Standardizing data followed Brodie et al. (1991) in prey categorization. Prey species were grouped into 3 categories based on Raw data were compiled and tabulated as fre- their known antipredator strategies: (1)