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Introduced European as main prey of the native carnivore (Lycalopex culpaeus) in disturbed ecosystems of central

Article in Studies on Neotropical Fauna and Environment · August 2013 DOI: 10.1080/01650521.2013.831521

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André V. Rubio Romina Alvarado University of Chile Agroadvance Ltda

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Studies on Neotropical Fauna and Environment Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/nnfe20 Introduced European rabbit as main prey of the native carnivore culpeo fox (Lycalopex culpaeus) in disturbed ecosystems of central Chile André V. Rubioab, Romina Alvaradoa & Cristian Bonacica a Laboratorio Fauna Australis, Departamento de Ecosistemas y Medio Ambiente, Facultad de Agronomía e Ingeniería Forestal, Pontificia Universidad Católica de Chile, Santiago, Chile b Facultad de Medicina Veterinaria y Zootecnia, Departamento de Etología, Fauna Silvestre y Animales de Laboratorio, Universidad Nacional Autónoma de México, México, D.F., México Published online: 12 Sep 2013.

To cite this article: André V. Rubio, Romina Alvarado & Cristian Bonacic (2013) Introduced European rabbit as main prey of the native carnivore culpeo fox (Lycalopex culpaeus) in disturbed ecosystems of central Chile, Studies on Neotropical Fauna and Environment, 48:2, 89-94, DOI: 10.1080/01650521.2013.831521 To link to this article: http://dx.doi.org/10.1080/01650521.2013.831521

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ORIGINAL ARTICLE Introduced European rabbit as main prey of the native carnivore culpeo fox (Lycalopex culpaeus) in disturbed ecosystems of central Chile André V. Rubioa,b*, Romina Alvaradoa & Cristian Bonacica aLaboratorio Fauna Australis, Departamento de Ecosistemas y Medio Ambiente, Facultad de Agronomía e Ingeniería Forestal, Pontificia Universidad Católica de Chile, Santiago, Chile; bFacultad de Medicina Veterinaria y Zootecnia, Departamento de Etología, Fauna Silvestre y Animales de Laboratorio, Universidad Nacional Autónoma de México, México, D.F., México (Received 30 March 2012; accepted 1 August 2013)

This study describes food habits of the culpeo fox (Lycalopex culpaeus) by scat analysis in two hills considered Priority Conservation Sites of the Mediterranean Region of central Chile, one of the most disturbed and modified ecosystems in the country. The results showed that from both sites consume the introduced European rabbit (Oryctolagus cuniculus) as primary prey and native small as secondary prey. This is the first record of feeding habits of the culpeo fox in central Chile that reports a higher consumption of European rabbit over pooled native small mammals. The reason for this result, as well as the higher trophic niche breadth and trophic niche diversity found in culpeo fox populations from the less disturbed hill, might be due to anthropogenic impacts on the study sites. However, to validate this claim, further studies on prey availability and human impact are required. The culpeo fox can have an important role in controlling pests such as the European rabbit; therefore, a more extensive study of this issue can bring important information for the conservation and management of natural areas of central Chile that show high human impact. Este estudio describe los hábitos alimentarios del zorro culpeo (Lycalopex culpaeus) en dos cerros considerados “Sitios Prioritarios de Conservación” de la eco-región mediterránea de Chile central, el cual es una de las zonas más intervenidas y modificadas del país. Los resultados encontrados muestran que los zorros de ambos cerros consumen como presa primaria al conejo europeo (Oryctolagus cuniculus) y a los pequeños mamíferos nativos como presa secundaria. Este es el primer registro de la dieta del zorro culpeo en Chile central que reporta un mayor consumo de conejo europeo por sobre el consumo de pequeños mamíferos nativos en su conjunto. La razón de este resultado, además de la mayor amplitud y diversidad de nicho trófico encontrado en la población de zorro culpeo en el cerro menos intervenido, podría deberse al impacto antropogénico en los sitios de estudios. Sin embargo, para validar esta afirmación se necesitan estudios sobre disponibilidad de presas e impacto humano. El zorro culpeo puede tener un papel importante en el control de plagas como el conejo europeo, por lo cual se requiere un estudio más acabado de esta temática, lo cual puede brindar información importante para la conservación y manejo de áreas naturales de Chile central que presenten un alto impacto antropogénico. Keywords: Chile; diet of carnivores; Oryctolagus cuniculus; sclerophyllous shrubland

Introduction et al. 1980, 1981; Simonetti 1986; Iriarte et al. 1989; The culpeo fox (Lycalopex culpaeus) is distributed in Ebensperger et al. 1991). The first studies of the culpeo fox’s diet, conducted around 35 ago, concluded

Downloaded by [UNAM Ciudad Universitaria] at 10:33 26 November 2013 South America from Colombia to (Jiménez & Novaro 2004). This wild canid is the that the relatively low predation of European rabbit largest of the Lycalopex and is considered by culpeo foxes and other endemic predators, such as the most carnivorous of all South American foxes the chilla fox (Lycalopex griseus) and Black-chested (Redford & Eisenberg 1992). Its diet ranges from wild Buzzard-eagle (Geranoaetus melanoleucus), was a sign ungulates to , depending on habitats and distri- that native predators had not yet learned to hunt butions (Jiménez & Novaro 2004). In sclerophyllous for this (Jaksic & Soriguer 1981). shrubland of central Chile, several studies on feeding European were introduced to Chile after habits through scat analysis report that culpeo foxes 1884 at several times, and in several parts of the prey mainly on native small mammals and secondarily country (Jaksic et al. 2002). on introduced European rabbits (Oryctolagus cunicu- Nowadays, in the sclerophyllous shrubland of the lus), in addition to other minor diet components, Mediterranean ecosystem of central Chile, a Global such as , , insects and fruits (e.g. Jaksic Biodiversity Hotspot (Myers et al. 2000), urbanization

*Corresponding author. Email: [email protected]

© 2013 Taylor & Francis 90 A. V. Rubio et al.

and intensive agriculture leave native areas with vary- native vegetation association comprises 21.2% of the ing degrees of deterioration (Bonacic & Ibarra 2009; total area, disturbed sclerophyllous shrubland cov- Pavéz et al. 2010). Because European rabbits are com- ers 77.3% and 1.5% is “other human impact” (Rojas mon in disturbed sites of sclerophyllous shrubland et al. 2010). Original sclerophyllous shrubland com- (Holmgren et al. 2000) and given the increasing inci- prises mainly plant communities of matorral com- dence of European rabbits in the diet of some raptors posed by Puya berteroniana and Echinopsis chiloensis (G. melanoleucus and Parabuteo unicinctus; Pavéz et al. and sclerophyllous forest of Cryptocarya alba, Peumus 1992, 2010; Jiménez & Jaksic 1993) and in the diet boldus, Quillaja saponaria and Lithraea caustica. of the culpeo fox in central Chile (Simonetti 1986; Disturbed sclerophyllous shrubland comprises com- Iriarte et al. 1989), we wanted to assess the degree munities mainly of Acacia caven, Colliguaja odorifera, to which rabbits are preyed upon by the culpeo fox Baccharis spp., Retanilla trinervia and other shrub by studying the diet of this carnivore in two sites species. The class “other human impact” includes fruit of high anthropogenic impacted sclerophyllous shrub- tree plantations and areas without vegetation (Rojas land which have never been previously studied. Our et al. 2010). aim was to identify their primary prey in each site and compare any differences in their diet. In addition, we compared our observations with other research on the Sample collection and analysis diet of culpeo foxes in central Chile. We assessed feeding habits of foxes using scat analy- sis, a non-invasive method that is relatively reliable and widely used (Putman 1984; Corbett 1989; Reynolds & Aebischer 1991; García & Kittlein 2005; Bianchi et al. Materials and methods 2010). During three seasons in 2009 (summer, win- Study area ter and spring) and one season in 2010 (autumn), we The study area comprises two neighboring hills 4 km collected scats of culpeo fox opportunistically along apart: Chena Hill (CH) of 1188 ha (33◦ 35 S, 70◦ 44 roads and trails in both study sites. We identified scats W) and Lonquén Hill (LH) of 4296 ha (33◦ 41 S, 70◦ by morphology (Chame 2003; Muñoz-Pedreros 2008), 45 W), located in the Santiago Metropolitan Region, smell, associated footprints and place of deposition. in central Chile. Both hills are surrounded by a matrix The culpeo fox is the only fox and the largest car- of rural and urban settlements that include several nivore inhabiting both sites. Scats that could not be roads, walls and fences throughout the sites, leaving confidently attributed to foxes were discarded. The them effectively as “islands” of habitat for terres- scats were washed in a 0.5 mm sieve and dried for trial wildlife (CONAMA 2004; Roa & Bonacic 2010). 12 h at 60◦C. Prey species were identified from frag- Although the distance between hills is short for culpeo ments of bones, teeth, hairs, feathers and exoskeletons, foxes, which can move a linear distance of 7.6 km using identification keys (Reise 1973; Chehébar & in a 24-h period and with a reported home range Martín 1989) and reference specimens. To determine average of 3.7 km2 in north-central Chile (Salvatori if the sample size of scats was adequate, we estimated et al. 1999), it is very difficult for culpeo foxes completeness by species accumulation curves using to move between sites given the reasons explained the Species Diversity and Richness 4.1.2 computer above. Therefore, we considered both sites indepen- package (Seaby & Henderson 2006) with 50 repeti- dent. Chilean authorities considered LH and CH tions for each study site, and then adjusted to the linear Downloaded by [UNAM Ciudad Universitaria] at 10:33 26 November 2013 “Priority Conservation Sites” as important remnants dependence model and the Clench model (Soberón & of the Mediterranean biome of central Chile, charac- Llorente 1993). For this analysis, the prey item was terized by sclerophyllous shrubland within an urban considered as a “species”. We defined “prey item” as matrix and a valuable habitat for some endangered any taxon that could be identified in the scats. For species (CONAMA 2004). Nevertheless, both study some categories, this taxon was the species, but higher sites reflect high anthropogenic impact in the form taxonomic levels (genus or order) were used for oth- of deforestation, agricultural expansion, pres- ers (Varela et al. 2008). We determined the importance sure, and grazing by rabbits, among others. Action for of each prey item in the diet as frequency of occur- conservation and management just began a few years rence (FO), which is the number of scats in which a ago with small local projects (Roa & Bonacic 2010). given prey item was found/total number of scats, and According to Rojas et al. (2010), CH is more disturbed relative abundance (RA) which is the minimum num- than LH. Original native sclerophyllous shrubland ber of individuals of a given prey item recorded in in CH covers 7.4% of the total area, 76.9% is dis- all scats/sum of all individuals recorded for all prey turbed sclerophyllous shrubland and 15.7% is “other items. Both indices are widely used for diet analy- human impact” (see below). At LH, the original sis including South American foxes (e.g. García & Studies on Neotropical Fauna and Environment 91

Kittlein 2005;Zapataetal.2005;Varelaetal.2008; Table 2. Comparison of frequency of occurrence (FO) and Silva-Rodriguez et al. 2010). The latter was calculated relative abundance (RA) of prey items recorded in scats only with vertebrate prey to improve comparisons with of Lycalopex culpaeus between the two study sites in the Santiago Metropolitan region, central Chile in 2009 and other studies (see below). We used the Chi-square 2010. test to compare the frequency of occurrence of the main prey items between and within sites. Trophic Chena Hill Lonquén Hill niche breadth of each study site based on frequency Prey item FO % RA % FO % RA % of occurrence was calculated using the standardized Levins’ index (Bstd;Krebs1989). The Levins’ index formula is B = 1/( pi2), where pi is the proportion Abrocoma benetti – – 23.4 18.4 Octodon degus 9.1 7.5 11.3 9 ofrecordsineachpreyitem(i). The standardized for- Phyllotis darwini 1.6 1.4 – – mula is Bstd = (B –1)/(Bmax –1),whereBmax is the Not identified 6.6 5.5 6.1 4.8 total number of prey categories recognized. We cal- Small mammals 17.5 14.4 41 32.2 culated the trophic diversity on each site using the pooled Shannon–Wiener index (Krebs 1989): H = – pi × Lagomorphs Oryctolagus 90 74 76 60 log2pi,wherepi is the proportion of the frequency cuniculus of prey item (i). The diversity values obtained were Birds 8.3 6.8 2.2 5.4 compared between sites by Hutcheson’s test (Krebs Eggs 4.6 4.8 6.8 1.8 1989). Finally, we made a comparison between RA of Reptiles main preys found in this study and other studies of the Lizards (Liolaemus) – – 0.7 0.6 Invertebrates culpeo fox around the Santiago Metropolitan Region Coleoptera 0.8 1.5 of Chile, by two-proportion Z-tests. We use the RA Fruits 4.1 3 instead of FO because all other studies used RA of Total number of scats 121 132 vertebrate preys as index. ∗ Total prey items 152 174

Note: ∗The sum of all individuals recorded for all prey categories. Results We collected a total of 121 scats from CH and 132 from LH. In each site the scats were sampled in similar numbers in each of the four seasons. Sample was significantly higher than FO of native small size was assumed to be adequate because both sites mammals pooled together (CH: χ 2= 132.8, df = 1, had an acceptable level of estimated completeness p < 0.001; LH: χ 2= 34.5, df = 1, p < 0.001; Table 2). ranging from 90% to 100%, depending on the model Comparing between sites, FO of European rabbits (Table 1). in scats was significantly higher in CH than in LH The diet was predominantly composed of (χ 2= 7.13, df = 1, p < 0.01; Table 2) and FO of small European rabbits and native small mammals, which mammals was significantly higher in LH than in CH together represented 92.2% of relative abundance in (χ 2= 16.79, df = 1, p < 0.001; Table 2). Trophic niche LH and 88.4% of CH respectively (Table 2). Eighty breadth was higher in LH (Bstd = 0.202) than CH per cent of the skulls and teeth of rabbits in the (Bstd = 0.116), as was the trophic diversity (H= 1.33, scats came from adult rabbits. Less abundant prey for LH; H= 1.21, for CH; t = 13.85, df = 304, Downloaded by [UNAM Ciudad Universitaria] at 10:33 26 November 2013 items were birds and beetles, among others (Table 2). p < 0.001). In both study sites, FO of European rabbits in scats Figure 1 shows the results of past studies of the diet of the culpeo fox in sclerophyllous shrubland of the Table 1. Parameters and predictions of the species accu- Santiago Metropolitan Region. We took the two stud- mulation models for prey items recorded in scats of ies that reported the highest level of consumption of Lycalopex culpaeus at two study sites in the Santiago rabbits, Simonetti (1986) and Iriarte et al. (1989)(let- Metropolitan region, central Chile in 2009 and 2010. ters c and d, see details in Figure 1), as our reference Linear dependent points. In CH, RA of rabbit remains was significantly model Clench model higher than that reported in any of the other studies ≥ < Site N Ar2 %A r2 % (z 3.4, p 0.001, in both cases), while RA of rabbit remains in LH was significantly higher only than study Chena Hill 8 8.3 0.91 96 8.9 0.97 90 d(z= 4.6, p < 0.0001). RA of native small mammal Lonquén Hill 9 9 0.95 100 10 0.99 90 remains was significantly lower compared with studies Note: N = number of prey items, A = the asymptote of the accumu- candd(CH:z≥ – 8.1, p < 0.0001 in both cases; LH: lation curves, r2 = coefficient of determination, % = completeness. z ≥ – 4.8, p ≤ 0.01 in both cases). 92 A. V. Rubio et al.

Figure 1. Comparison of relative abundance (RA, in %) of native small mammals and European rabbits (Oryctolagus cuniculus) in scats of Lycalopex culpaeus between several studies performed in the Santiago Metropolitan District of Chile, and our results. Abbreviations: a. Jaksic et al. (1980); b. Jaksic et al. (1981); c. Simonetti (1986); d. Valley site, Iriarte et al. (1989); e. Mountain site, Iriarte et al. (1989); f. Ebensperger et al. (1991); g. Lonquén Hill, this study; h. Chena Hill, this study.

Discussion Sovada et al. 2001; Rau & Jiménez 2002; Pia et al. This study reported prey items commonly described in 2003; Farias & Kittlein 2008). In the sclerophyllous the literature of the culpeo fox’s diet in central Chile, shrubland of central Chile, human disturbance may where native rodents and the European rabbit com- favor the European rabbit and adversely affect native prise the main prey items, while birds and reptiles, small mammals (Pavéz et al. 2010). This is because among others, are minor components of their diet (e.g. rabbits use areas under shrub canopies as habitat, but Simonetti 1986; Iriarte et al. 1989; Ebensperger et al. they also use extensive open areas that are the result 1991). However, this is the first study that recorded of human impact (Simonetti 1989a; Holmgren et al. the European rabbit as the culpeo fox’s primary prey 2000), while native small mammals use mainly the species compared with the total native small mammals areas under shrub canopies as microhabitat and most in central Chile. Previous studies report higher fre- of them avoid open areas (Meserve 1981; Simonetti 1989b). In fact, the degu (Octodon degus) and the

Downloaded by [UNAM Ciudad Universitaria] at 10:33 26 November 2013 quencies of European rabbits than of any other species of native small mammals in the scats of culpeo foxes Bennett’s chinchilla rat (Abrocoma benetti), which are (Simonetti 1986; Iriarte et al. 1989), but they date back among the main native prey species of culpeo foxes in about 25 years, therefore new research on the trophic central Chile (Jaksic et al. 1981; Simonetti 1986; Iriarte ecology of culpeo foxes should be performed in the et al. 1989; Ebensperger et al. 1991), may be decreasing same sites that were studied in the 1980s (Simonetti in number due to anthropogenic disturbance in small 1986; Iriarte et al. 1989; Ebensperger et al. 1991). This fragments (Fernández & Simonetti 2013) and larger might help to assess possible variation in the feeding areas of sclerophyllous shrubland (Pavéz et al. 2010). behavior of L. culpaeus and to better understand the In this context, and given that the food habits of the importance of the European rabbits in their diet over culpeo fox can change according to prey availability time. (e.g. Johnson & Franklin 1994; Silva et al. 2005), the Anthropogenic landscape changes have been high frequency of rabbit remains and the lower fre- shown to increase spatial variability in carnivore quency of degus and Bennett’s chinchilla rat remains diets, even over small distances, by favoring newly reported in this study might be due to anthropogenic introduced species and altering the distribution and changes in native habitats. In addition, a more diverse abundance of native species (e.g. Fedriani et al. 2001; diet of the culpeo foxes at LH (less disturbed site) than Studies on Neotropical Fauna and Environment 93

at CH (most disturbed site) is probably due to dif- Corbett LK. 1989. Assessing the diet of dingoes from : a ferences in resource availability as a consequence of comparison of 3 methods. J Wildlife Manage. 53(2):343–346. anthropogenic impact. However, further research on Diuk-Wasser MA, Cassini MH. 1998. A study on the diet of the minor grisons and a preliminary analysis of their role in the prey availability and human disturbance in these sites control of rabbits in Patagonia. Stud Neotrop Fauna Environ. should be addressed to sustain this claim. Another 33(1):3–6. point to be considered, which requires further studies, Ebensperger L, Mella J, Simonetti JA. 1991. Trophic-niche relation- is the consumption of rabbits captured in snare-traps. ships among cuja, Dusicyon culpaeus, and Tyto alba in Given that usually adult rabbits are the most often central Chile. J . 72(4):820–823. Farias AA, Kittlein MJ. 2008. Small-scale spatial variability in the trapped with snares whereas juveniles are most fre- diet of foxes (Pseudalopex gymnocercus) and human- quently dug up from their warrens by culpeo foxes induced changes in prey base. Ecol Res. 23(3):543–550. (Jaksic et al. 1980), snare-robbing might explain the Fedriani JM, Fuller TK, Sauvajot RM. 2001. Does availability of high relative abundance of adults rabbits in the diet anthropogenic food enhance densities of omnivorous mammals? of culpeo foxes at Chena and Lonquén Hills, which An example with coyotes in southern California. Ecography. 24(3):325–331. are both sites where we observed high illegal hunting Fernández IC, Simonetti JA. 2013. Small mammal assemblages in activity with snare-traps. fragmented shrublands of urban areas of Central Chile. Urban We suggest that given the high rabbit predation Ecosyst. 16(2):377–387. by culpeo foxes, their potential role as regulators of García VB, Kittlein MJ. 2005. 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