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Predatory Impact of Rainbow Trout on Native Fish in the Cape Size Predatory impact of non-native rainbow trout on endemic fish populations in headwater streams in the Cape Floristic Region of South Africa Jeremy M. Shelton, Michael J. Samways & Jenny A. Day Biological Invasions ISSN 1387-3547 Biol Invasions DOI 10.1007/s10530-014-0735-9 1 23 Your article is protected by copyright and all rights are held exclusively by Springer International Publishing Switzerland. This e- offprint is for personal use only and shall not be self-archived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Biol Invasions DOI 10.1007/s10530-014-0735-9 ORIGINAL PAPER Predatory impact of non-native rainbow trout on endemic fish populations in headwater streams in the Cape Floristic Region of South Africa Jeremy M. Shelton • Michael J. Samways • Jenny A. Day Received: 27 September 2013 / Accepted: 3 June 2014 Ó Springer International Publishing Switzerland 2014 Abstract Non-native rainbow trout Oncorhynchus trout. The mean densities of native Breede River redfin mykiss have been widely introduced in the Cape Pseudobarbus burchelli, Cape kurper Sandelia capen- Floristic Region (CFR) at the south-western tip of sis and Cape galaxias Galaxias zebratus, were 89–97 % Africa and may pose a serious threat to endemic lower in invaded streams than in streams without trout. freshwater fishes in the region. Quantitative informa- Furthermore, while native fish were present at all 12 tion about trout impacts in the CFR is scarce but is sites without trout, they were not recorded at seven of urgently needed to guide legislation and management the 12 invaded sites. None of the measured environ- efforts. We used a combination of comparative and mental variables differed significantly between sites experimental approaches to evaluate impacts of intro- with and without trout, and distance-based linear duced trout on native fish populations in headwater models identified trout density as the best predictor of streams draining the upper Breede River catchment in redfin and kurper density, while galaxias density was the CFR. Fish populations were surveyed, and 19 best predicted by other environmental variables (ripar- environmental variables measured, in 24 minimally- ian vegetation, canopy cover, substrate length, site disturbed streams, half of which had been invaded by slope). Native fish B40 mm in length were largely absent from invaded streams, but generally abundant in streams without trout, and a field experiment confirmed Electronic supplementary material The online version of that trout selectively consume small redfin. Taken this article (doi:10.1007/s10530-014-0735-9) contains supple- mentary material, which is available to authorized users. together, these findings constitute evidence that trout have depleted the abundance of CFR-endemic fishes J. M. Shelton (&) Á J. A. Day through size-selective predation. It is recommended University of Cape Town, that managers aim to prevent new trout introductions Private Bag X3, Rondebosch, Cape Town 7701, South Africa and consider eradicating trout populations where they e-mail: [email protected] pose a threat to highly threatened native species. J. M. Shelton Keywords Invasive Á Oncorhynchus mykiss Á DST-NRF Centre of Excellence for Invasion Biology (CIB), Private Bag X1, Matieland, Stellenbosch Predation Á Threatened native fish 7602, South Africa Introduction M. J. Samways Stellenbosch University, Private Bag X1, Matieland, Stellenbosch 7602, The rainbow trout Oncorhynchus mykiss (Walbaum South Africa 1792) is among the most widely introduced fish in the 123 Author's personal copy J. M. Shelton et al. world (Fausch 2007). From its native range in Pacific endemic to the region (Skelton 2001; Chakona and North America and eastern Russia, it has been Swartz 2013), but nearly all of these endemic species introduced to at least 97 countries, and to every appear in the categories vulnerable, endangered and continent except Antarctica (Crawford and Muir critically endangered of the IUCN Red List (IUCN 2013). 2008). Rainbow trout are gape-limited predators Threats include habitat loss and fragmentation, hydro- (Lescak and von Hippel 2011), and while as juveniles logical alteration, climate change, overfishing and pollu- they feed mostly on small invertebrates, they can tion, but the greatest threat is that posed by introduced consume increasingly larger prey, such as amphibians predatory fish (Tweddle et al. 2009; Chakona and Swartz and fish, as they grow (Raleigh et al. 1984). In many 2012). To date, 16 species of non-native freshwater fish cases, introduced rainbow trout have established self- have established self-sustaining populations in the CFR sustaining populations (Fausch 2007) and negative (Marr 2012), and several of these species have been impacts on native fish populations, largely as a result linked to declines in native fish populations (de Moor and of predation, have been reported from across the globe Bruton 1988;Tweddleetal.2009). [e.g. New Zealand (McDowall 2006), Australia Interactions with non-native fish, combined with (Crowl et al. 1992; Lintermans 2000; McDowall the effects of habitat degradation, have led to the 2006), South America (Young et al. 2010; Habit et al. disappearance of native fish from the middle and lower 2010), North America (Dunham et al. 2004) and in sections of many rivers in the CFR (Tweddle et al. Japan (Kitano 2004)]. Because of their widespread and 2009; de Moor and Day 2013; Weyl et al. 2013). severe impacts, rainbow trout have been listed by the Consequently, the region’s native fish populations World Conservation Union Global Invasive Species have become highly fragmented, and many species are Programme (GISP) as one of the World’s 100 Worst now largely restricted to headwater reaches of streams Alien Invasive Species (Lowe et al. 2000). Although (Swartz et al. 2004; Tweddle et al. 2009; Chakona and the consequences of rainbow trout introductions have Swartz 2012). Headwater streams have generally been received detailed study in many areas, impacts in other less severely impacted by human-related activities locations, such as South Africa, remain largely than lower-lying reaches because they are situated in unknown. mountainous areas that are difficult to access and Rainbow trout (henceforth ‘‘trout’’) were intro- unsuitable for agriculture, human settlements and duced to South Africa in 1897 for angling purposes (de reservoirs (Swartz et al. 2004; Tweddle et al. 2009;de Moor and Bruton 1988), and widespread stocking of Moor and Day 2013). Consequently, the headwater the country’s rivers ensued (Hey 1926). Today trout reaches of streams function as native fish refugia occur in *75 % of South Africa’s major river systems, within the region’s highly degraded riverscapes, and and while self-sustaining populations are generally are thus critical habitats from a conservation perspec- limited to the cooler headwater reaches of streams, tive. Trout are well adapted to environmental condi- many other populations are sustained through contin- tions in these headwater streams (Raleigh et al. 1984) ual stocking (van Rensburg et al. 2011). Beginning in and may pose a serious threat to remaining native fish the 1960s, conservation organizations gradually populations in the CFR (Cambray 2003). became aware that trout appeared to be having a To our knowledge, there has only been one previous negative impact on native aquatic species, particularly assessment of trout impacts on native fish in the CFR; fishes (de Moor and Bruton 1988). Despite the fact that that of Woodford and Impson (2004), who studied negative impacts of trout in South Africa have long interactions between trout and three species of native been suspected, evidence is mostly circumstantial and fish and found some evidence for spatial segregation anecdotal (de Moor and Bruton 1988; Cambray 2003). between trout and native fish and predation by trout on Quantitative information on trout impacts in South one native species, the Cape galaxias Galaxias zebratus Africa is urgently needed to guide legislation and (Castelnau 1861). Although their study was confined to management efforts (Cambray 2003). five pools within a single stream (the upper Berg River), The Cape Floristic Region (CFR) at the south-western their findings are consistent with the view that trout may tip of South Africa is a hotspot for freshwater fish have a negative predatory impact on native fish endemism (Linder et al. 2010; de Moor and Day 2013). populations in CFR streams. The aim of the present Of the 20 species that have been described, 17 are study was to expand on their preliminary work by 123 Author's personal copy Predatory impact of non-native rainbow trout on endemic fish populations Fig. 1 Location of sampling sites in the upper Breede River with trout. The numbers of the sampling sites correspond to the catchment in the CFR of South Africa. White circles represent numbers in Supplementary Material 1. Names of towns, as well sampling sites without trout, and black circles represent sites as major rivers and reservoirs, are shown evaluating the predatory impact of introduced trout on tributaries of the Sonderend River upstream of Thee- CFR-endemic fishes through a combination of compar- waterskloof Reservoir (Fig. 1). The area experiences a ative and experimental approaches. We surveyed fish Mediterranean climate (Cowling and Holmes 1992)and populations, and measured environmental variables, at has a mean annual rainfall of *800 mm (Steynor et al. sites on 24 headwater streams in a catchment partially 2009). Natural vegetation is predominantly Sandstone invaded by trout to assess their impacts on native fish Fynbos (Rebelo et al.
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