Distribution of Non-Native Brook Trout (Salvelinus Fontinalis) Across Norwegian Waterbodies – Is It an Invasive Species?
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Fauna norvegica 2018 Vol. 38: 1-8 Distribution of non-native brook trout (Salvelinus fontinalis) across Norwegian waterbodies – is it an invasive species? Trygve Hesthagen1, Geir H. Bolstad1 and Einar Kleiven2 Hesthagen T, Bolstad GH and Kleiven E. 2018. Distribution of non-native brook trout (Salvelinus fontinalis) across Norwegian waterbodies – is it an invasive species? Fauna norvegica 38: 1-8. Non-native brook trout (Salvelinus fontinalis MITCHILL, 1815) was introduced into Norway in 1883. However, it was not until the late 1970s that this acid-tolerant salmonid species was stocked into many acidified lakes that many populations became established. In 2004, all brook trout stocking in Norway ceased. In this study, we surveyed the distribution of brook trout in Norwegian water bodies. A totally of 202 self-sustaining populations were identified, mostly in unregulated lakes (n=101), streams (n=71) and also to some extent in reservoirs (n=25). Only four populations were found in inland rivers, and one population in a river with Atlantic salmon (Salmo salar L.). Localities with brook trout covered a wide range of altitudes and sizes. Analyses of time-series catches in sympatric populations of brown trout (Salmo trutta L.) and brook trout from 12 lakes (1997-2012) revealed a strong decline in brook trout stocks. A similar development in abundance has also emerged from other studies in recent years. Generally, lake-dwelling brook trout is regarded as a low-risk species with respect to invasiveness. However, brook trout/brown trout interactions may be habitat-specific, as brook trout may dominate in small and relatively cold streams. doi: 10.5324/fn.v38i0.2255. Received: 2017-04-04. Accepted: 2017-12-05. Published online: 2018-01-17. ISSN: 1891-5396 (electronic). Keywords: Streams, lakes, brook trout, non-native, invasiveness. 1. Norwegian Institute for Nature Research (NINA), PO Box 5685, NO-7485 Trondheim. 2. Norwegian Institute for Water Research (NIVA), Jon Lilletuns vei 3, NO-4879 Grimstad, Norway. Corresponding author: Trygve Hesthagen E-mail: [email protected] INTRODUCTION of freshwater fishes have been carried out over the 19th century (Gozlan 2008; Clavero & Villero 2013; Gozlan et al. Invasive species, i.e. non-native species that spread beyond 2010). These have included three salmonid species; brook the introduction site and become abundant and established trout (Salvelinus fontinalis MITCHILL, 1815), rainbow trout (Rejmanek et al. 2002), are increasingly recognized as one of (Oncorhynchus mykiss WALBAUM, 1792) and brown trout the main threats to biodiversity in fish communities at a gobal (Salmo trutta L., 1758) (MacCrimmon & Marshall 1968; level (García-Berthou 2007; Trochine et al. 2017). Freshwater MacCrimmon & Campbell 1969; MacCrimmonn 1971; Scott & ecosystems are particularly affected by invasive fish species Crossman 1973). Brook trout are native to northeastern North (Clavero & Garcia-Berthou 2006), which have become a America (MacCrimmon & Campbell 1969; Scott & Crossman growing source of concern worldwide for several decades 1973). However, since the late 1800s, attempts have been made (Kohler & Courtenay 1986; Sala et al. 2010; Gallardo & Aldridge to establish brook trout all over the world, mainly due to its 2015). The management of invasive species is therefore one of appeal as a sport fish. the major challenges that conservation biologists will face in the Brook trout were successfully introduced into Norwegian near future (Allendorf & Lundquist 2003). waters in 1883 (Hesthagen & Kleiven 2013; Kleiven & Hesthagen Several large-scale intercontinental introductions 2013). However, subsequent releases of fry at 20-30 locations 1 Hesthagen, Bolstad and Kleiven: Brook trout in Norway Fauna norvegica 38: 1-8. 2018 produced only one known self-sustaining population, in METHODS Overnbekken stream in Buskerud County (Eken 1988). In 1917 or 1918, another population of unknown origin was established The distribution of brook trout in Norwegian water bodies was from fry stocking in Øyfjell, Telemark County (Huitfeldt-Kaas surveyed in 2012-2016 (Hesthagen & Kleiven 2013; Hesthagen 1924). In this area, brook trout were since translocated to several & Sandlund 2016). Data was obtained via literature studies and sites (Grande 1964; Hesthagen & Kleiven 2013). In the late interviews with fishermen, landowners, fishery organizations, 1970s, large-scale stocking of brook trout started in southern and local and state environmental authorities. First, we grouped Norway. This mitigation measure was intended to compensate their occurrence at county level, and thereafter by individual for the extensive damage to fish caused by acidification, as watersheds. On both scales, possible occurrence of brook trout more than 9600 populations, mainly of brown trout, became was identified, starting at the highest lake in each watershed. extinct (Sevaldrud & Muniz 1980; Hesthagen et al. 1999). As Downstream localities were then surveyed in order to establish brown trout in most cases was the only native fish species, large the existence of brook trout. Their longevity is usually less than areas became completely devoid of fish. The reason for stocking four years in Norwegian waters (Qvenild 1986; Sægrov et al. brook trout was that it was recently recognized as an extremely 2008). As stocking ceased in 2004, all populations that were acid-tolerant species (Dunson & Martin 1973; Trojnar 1977; identified in 2012 at the latest are considered self-sustaining. Muniz & Leivestad 1979). At least 12 hatcheries in southern Each stock was categorized according to the type of habitat Norway have produced offspring (age 0+) of brook trout in it occupied; (i) unregulated lakes, (ii) regulated lakes [reservoirs recent decades (Hesthagen & Kleiven 2014). Most lakes were for hydropower generation], (iii) first- and second-order inland probably stocked with at least several hundred individuals. In streams, (iv) rivers with a maximum width of 4-5 m, and (v) regulated lakes, however, compensatory stockings were carried rivers with anadromous salmonids. A stock of brook trout is out by the power companies, and in most cases these amounted defined as one that inhabits a single lake, river or stream. Any to between 1 000 and 10 000 individuals annually (Møkkelgjerd given lake is defined as containing only one stock, irrespective & Gunnerød 1985). of whether spawning could occur in several tributary streams. In 2004, the stocking of brook trout in Norwegian waters Streams up- or downstream of lakes containing brook trout, or was stopped (Hesthagen & Sandlund 2007). The environmental streams between two lakes with brook trout, are not regarded authorities realized that introductions of this non-native species as separate stocks. Nor are stocks in tributaries or second-order were at odds with the internationally adopted goal of conserving inland rivers, if the first-order stream or river contains brook native biodiversity. Moreover, brown trout populations in trout. However, stocks in tributaries to a river with Atlantic previously acidified areas had recently started to recover salmon (Salmo salar L.) are defined as reproductive units if that (Hesthagen et al. 2011a, 2016). This is thanks to improved river does not host brook trout. We used different categories for water quality through a substantial reduction in sulphates in lake size and altitude (both streams and lakes). precipitation (cf. Garmo et al. 2014). In addition, large scale We analyzed the change in relative abundance of brook liming campaigns have been carried out since the 1980s trout living in sympatry with brown trout from 11 un-limed (Sandøy & Romundstad 1995). lakes in upper stretches of the Tovdal watershed (170 to 218 When a non-native fish species is introduced into a body m a.s.l.), Aust-Agder County, and from one limed lake in Otra of water where it did not previously occur, there are several watershed, Lake Dåsvatn at 190 m a.s.l., Vest-Agder County, possible outcomes (Gozlan et al. 2010; Blackburn et al. 2011; based on catches between 1997 and 2012. Only these two Gallardo & Aldridge 2015). The negative ecological impacts species of fish are found in these 12 lakes. The lakes in Tovdal vary highly depending on the species involved, community watershed were sampled only with gillnets with mesh sizes structure and abiotic environmental conditions (Savini et al. between 15-24 mm (knot to knot), from April to August every 2010; Henriksson 2016 a, b). For the majority of non-native year. Sampling took place several times each month, using species, no quantitative information is available regarding the between 40 and 100 gillnets. Not all lakes were sampled every consequences of such introductions (Simberloff et al. 2013). year in Tovdal watershed. In Lake Dåsvatn, sampling was Brook trout is classified as a medium-risk species among non- mainly carried out with traps with mesh size of 10 mm, between native freshwater fish (Copp et al. 2009; Britton et al. 2010). Our mid-April and late November. Gillnets with 19.5 to 24 mm mesh article describes the distribution of brook trout in Norwegian sizes were also occasionally used. The total catch of brown trout water bodies. Furthermore, we studied their development in and brook trout in lakes in the Tovdal and Otra watersheds were recent years in 12 lakes, based on time-series. The specific 334251 vs. 1634 and 45162 vs. 925 individuals, respectively. objective of the study is to assess whether brook trout is an These catch data from Lake Dåsvatn were analyzed using a invasive species. mixed effect model with a logit link and binomially distributed residuals: 2 Hesthagen, Bolstad and Kleiven: Brook trout in Norway Fauna norvegica 38: 1-8. 2018 E[Logit(yj)] = b0 + b1(Year j – 1997) + cj RESULTS where E[Logit(yj)] is the expected proportion of brook trout A total of 202 self-sustaining populations of brook trout were in the catch (on logit scale) for each year j, b0 is the intercept, recorded in southern Norway, mainly in the counties of Aust- b1 the slope, and c a random factor distributed according to an Agder (n=62), Vest-Agder (n=49) and Telemark (n=38) (Figure AR(1) process to account for the non-independence among the 1).