Coexistence Between the Southern River Otter and the Alien Invasive North American Mink in Marine Habitats of Southern Chile G
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bs_bs_bannerJournal of Zoology Journal of Zoology. Print ISSN 0952-8369 Coexistence between the southern river otter and the alien invasive North American mink in marine habitats of southern Chile G. Medina-Vogel1, M. Barros1, J. F. Organ2 & L. Bonesi3 1 Facultad de Ecología y Recursos Naturales, Universidad Andres Bello, Santiago, Chile 2 U.S. Fish and Wildlife Service, Hadley, MA, USA 3 C/O Department of Earth Sciences, University of Cambridge, Cambridge, UK Keywords Abstract competition; diet; habitat; alien mink; otter. The interaction between native and introduced predators can be an important Correspondence determinant of the success of introduced species and of the magnitude of their Gonzalo Medina-Vogel, Universidad Andres effects. In Europe, it has been shown that the American mink Neovison vison can Bello, República 252. Santiago, Chile. be affected by native competitors; however, such evidence has not been found in Email: [email protected] South America. We studied the southern river otter Lontra provocax and the American mink at five marine sites, where they coexisted, and at one freshwater Editor: Andrew Kitchener site, where only mink were present, in southern Chile. We used the signs of both species to study their habitat use and diet, and radio tracking to study their activity Received 10 April 2012; revised 12 patterns. The results indicated that otters and minks tended to use different November 2012; accepted 20 November habitats in marine environments, the otter favouring littoral areas that are rocky 2012 and steep while the mink favours areas of gravel with a gently sloping intertidal zone. These differences were also reflected in their diets. At one of the coastal sites doi:10.1111/jzo.12010 where the diet of the two species was similar, the activity pattern of minks was mostly diurnal, which is unusual. While differential habitat use may be the way through which the American mink is able to coexist with the southern river otter in coastal habitats of southern Chile, it is possible that otters are having an effect on individual minks by affecting their activity patterns, although more observa- tions are needed to confirm this hypothesis. Introduction Nearctic region, has been introduced throughout the world including southern Chile where it is now present both in Intraguild predation, the killing of species that use similar freshwater and marine habitats (Medina, 1997). In coastal resources and the non-lethal effects that can ensue (Cress- habitats of southern Chile, the American mink coexists with well, 2008), can strongly determine the behaviour, condi- the southern river otter Lontra provocax, an endangered tions, population density and distribution of animals species that is slightly larger than the Eurasian otter (about (Palomares & Caro, 1999; Sergio & Hiraldo, 2008), and its 10–20%) (Sepúlveda et al., 2007). As the effects of intraguild effects are common in carnivorans (Arim & Marquet, 2004). predation are strongly dependent on size differences between Eurasian otters Lutra lutra are known to attack and kill predator and prey (Palomares & Caro, 1999), we expect the American mink Neovison vison (Bonesi & Macdonald, 2004a; southern river otter to have similar if not greater effects on Simpson, 2006), and a number of studies have detected the American mink than the Eurasian otter. In particular, at effects of otters on mink both at the population and at the a local scale, if the larger southern river otter poses a threat individual level. For example, Eurasian otters can affect the to the mink, the mink should avoid close contact or intense mink’s distribution and population density in the United site-use overlap (Tannerfeldt, Elmhagen & Angerbjörn, Kingdom (Bonesi & Macdonald, 2004a; Bonesi, Strachan & 2002). To test for the possible effects of the southern river Macdonald, 2006; Mcdonald, O’Hara & Morrish, 2007), otter on the American mink, we focused on the following induce a shift in their diet (Clode & Macdonald, 1995; questions: (1) Is the presence of American mink negatively Bonesi, Chanin & Macdonald, 2004; Harrington, Harrington correlated to that of the southern river otter at sites where & Macdonald, 2009) and change their circadian activity both species are present?; (2) What role does habitat play in patterns (Harrington et al., 2009), although these effects the coexistence of the two competitors and is this reflected in are not always observed (Brzezin´ski, S´wie˛cicka-Mazan & their diets?; (3) Do mink show a tendency to be more diurnal Romanowski, 2008). The American mink, a native of the than expected when they coexist with otters? Journal of Zoology •• (2013) ••–•• © 2013 The Zoological Society of London 1 Coexistence between introduced mink and native otter G. Medina-Vogel et al. Materials and methods and located between the low-tide line and 50 m inland above the high-tide line. The average width of the intertidal zone Six study sites, five marine and one freshwater, were randomly varied according to the seashore gradient (range 3–50 m). The selected between 44°30’S–72°34’W and 44°51’S–72°56’W in habitat characteristics that were recorded are listed in Table 1. the southern Chilean archipelago (Fig. 1). Each site was 3 km At marine sites, vegetation distance was measured as the dis- long and separated by more than 40 km from neighbouring tance between where vegetation started above the intertidal sites for independence (Niemimaa, 1995; Sepúlveda et al., zone and the high-tide line. At the freshwater site, this distance 2007). The study area is covered by abundant vegetation was calculated from the interface of the water surface and dominated by temperate Norpatagonicus Valdivian rainforest river bank to where the vegetation started. Substrata were (Veblen & Schlegel, 1982), the average annual temperature is measured in the intertidal area and classified based on the 9°C, and up to 2973 mm of rainfalls annually. There is only diameter of their components as: sand (<0.5 cm), gravel (0.5– low-impact human activity along the entire study site as indi- 10.0 cm), small rocks (10.0–25.0 cm) and large rocks (>6 m). cated by fishing boats and a few scattered houses. All six study sites (marine and freshwater) were surveyed every 3 months between May 2007 and February 2009 for a total of five to seven visits depending on weather conditions, Habitat use and overlap and between 10 and 15 days were needed to complete each Habitat characteristics of the marine study sites were assessed survey session. Each study site was surveyed along its shore- using 10-m-wide transects spaced at 100 m from each other line either by boat or on foot, and mink and otter field signs CHILE 73°37W ARGENTINA QF2 MF QF1 Figure 1 Geographical locations of marine and river study sites. The sites names are: Magdalena Island MI Magdalena Fiord (MF); Queulat Fjord (QF1); 44°58S Queulat Fjord (QF2); Magdalena Island (MI); CR MV Marta Valley (MV) – all marine habitats; and Cisne River (CR). Table 1 Results of the survey of the habitat characteristics for the marine (MF, QF1, QF2, MI, MV) and the freshwater (CR) sites Site name Orientation Exposure to waves Maximum vegetation distance Intertidal slope Substratuma MF South Protected 30 m 60% 15°–30° 60% Gravel 40% > 80° 40% Large rocks QF1 North Protected 10 m 100% > 80° 100% Large rocks QF2 North Protected 10 m 100% > 80° 100% Large rocks MI East Protected 30 m 100% < 15° 100% Gravel MV North Exposedb 10 m 100% < 15° 60% Sand 40% Gravel CR South – 30 mc 100% > 60°c 100% Small rocks aIn the marine study sites, substratum refers to the kind of material present in the intertidal area, while in the freshwater site (CR) substratum was measured between the water level and the vegetation. bExposed to winter north winds. cVegetation cover and slope considered from or above water level. CR, Cisne River; MF, Magdalena Fjord; MI, Magdalena Island; MV, Marta Valley; QF1, Queulat Fjord 1; QF2, Queulat Fjord 2. 2 Journal of Zoology •• (2013) ••–•• © 2013 The Zoological Society of London G. Medina-Vogel et al. Coexistence between introduced mink and native otter (footprints and faeces) were searched for. One month before Diet the start of the study in 2007, a preliminary survey was carried Spraints were collected at all six study sites during the surveys out to map otter and mink latrines (later call stations), and to described earlier. They were washed and dried at 75°C for collect and eliminate field signs – these samples were not used 48 hours (Medina, 1997). Prey remains were identified and in the analyses. compared with reference material at the Ecosystem Health Habitat use and habitat overlap between the two species Laboratory, Universidad Andrés Bello. Diet assessment was were assessed by using otter and mink field signs as a proxy for carried out by prey species and prey groups (fish, crustaceans, their presence and by applying the index of revisitation rate terrestrial and other aquatic species). The results were tabu- (IRR) method that was developed and tested in previous lated as: ‘O,’ occurrence (number of spraints in which a prey studies (Medina-Vogel et al., 2003; Medina-Vogel & species or prey group occurred) and ‘RF,’ relative frequency Gonzalez-Lagos, 2007). Surveys for mink signs have been (number of spraints in which a prey species or prey group shown to be a sufficiently reliable method to detect the occurred divided by the total occurrence of all the prey species presence/absence of mink even when otters are present (Bonesi or prey groups found). The data were not normally distributed & Macdonald, 2004b; Harrington, Harrington & Macdonald, even after transformation. Therefore, differences between 2008). Moreover, an experimental study of the reaction of prey groups were assessed by means of non-parametric tests mink to Eurasian otters’ odour has shown that mink do not (Pearson chi-square test, Kruskal–Wallis one-way analysis of avoid otter markings (Harrington et al., 2009).