A Look Into Salinity Tolerance of the Round Goby
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A look into salinity tolerance of the round Goby: A tale of two salinities Felix Englund Örn Degree project for Master of Science in Biology Animal Ecology, 60 hec, vt 2017 Department of Biological and Environmental Sciences University of Gothenburg Supervisor: Charlotta Kvarnemo 1 Examiner: Staffan Andersson Table of contents Abstract ................................................................................................................................................. 3 Introduction ................................................................................................................................................... 3 Aims & Predictions ........................................................................................................................................ 6 Method .......................................................................................................................................................... 7 Fish sampling, upkeep and acclimatization ........................................................................................... 7 Figure 1. ................................................................................................................................................. 8 Table 1. .................................................................................................................................................. 9 Table 2 ................................................................................................................................................. 10 Figure 2 ................................................................................................................................................ 11 Behavioral Experiment 1: Activity ....................................................................................................... 11 Behavioral Experiment 2: Salinity Choice ............................................................................................ 11 Figure 3. ............................................................................................................................................... 12 Statistics & null hypothesis .................................................................................................................. 12 Figure 4. ............................................................................................................................................... 13 DNA Extractions ................................................................................................................................... 14 DNA Concentrations ............................................................................................................................ 14 Results ..................................................................................................................................................... 15 Behavioral Experiment 1: Activity ....................................................................................................... 15 Figure 5 ................................................................................................................................................ 15 Behavioral Experiment 2: Salinity Choice ............................................................................................ 15 Figure 6. ............................................................................................................................................... 16 Discussion ................................................................................................................................................ 17 Figure 7. ............................................................................................................................................... 16 Figure 8. ............................................................................................................................................... 17 Acknowledgements ............................................................................................................................. 20 References ................................................................................................................................................... 21 2 Abstract The round goby is a highly successful invasive species of benthic fish. Transport via ballast water has led to its establishment in the Baltic Sea, large harbors and rivers across Europe and the Laurentian Lakes of North America. Generalist behavior and wide tolerance to abiotic factors enables it to thrive in many environments and recent studies show salinity tolerance to be higher than previously reported, with suggestions that tolerance is linked to the salinity of origin. Establishment of round gobies in Gothenburg, Sweden, has provided a natural experiment with exposure, for the first time, to almost oceanic conditions. The aim of this study was to investigate Gothenburg round goby populations at two different salinities in order to reveal if variation in salinity tolerance is due to genotypic sorting of genetically pre-adapted individuals, or to phenotypic plasticity. This was addressed using behavioral activity experiments which find that round gobies are equally active across point of capture, suggesting that salinity tolerance is due to phenotypic plasticity. Importantly, the presence of a wild population, apparently unaffected by near oceanic conditions, suggests that high salinity is not a natural constraint on dispersal in the round goby. Furthermore, local findings indicate intraspecific competition may drive the fish to actively seek unpopulated areas with higher salinities. as an invader stems in part from a generalist Introduction feeding behavior (Nurkse et al., 2015), multiple spawning’s per breeding season The round goby (Neogobius melanostomus) (Meunier et al., 2009), an ability to tolerate a is a species of benthic fish native to the wide range of salinities and temperatures as Caspian and Black seas. Through well as a tolerance to low oxygen levels, anthropogenic activities it has spread far and which enables it to thrive in many wide across Europe as well as to the Great environments (Kornis et al., 2012). Lakes of North America (Brown and Stepien, Throughout its history as an invader it has 2008) and has established itself as one of the established in a wide range of aquatic widest spread invasive fish species in the environments. Round goby populations have world (Kornis, 2012). As with any invasive previously been found in fresh to brackish species, it is of great concern how the native water (0 - approx. 15‰) and the species has ecosystem will acclimate to its introduction established itself in large European rivers and how future management should be (e.g. Rhine and Elbe) and in the Laurentian conducted. The round goby has been shown lakes of North America (freshwater) and the to affect ecosystems both negatively and Baltic Sea (brackish), but no populations positively and it has proven to be a complex have been observed in full oceanic matter affecting multiple trophic levels conditions (Charlebois et al., 2001; Kornis, et (Hirsch, 2015; Kornis et al., 2012). Its success al. 2012). However, a tolerance of up to 40‰ 3 has been reported (Kullander et al. 2012), An establishment in a new although Kornis et al. (2012) noted that environment, with new selective pressures, these results derive from the Caspian Sea such as high salinity, sets the stage for local and Aral Sea. In these oceans CaSO4 is the adaptation. Local adaptation, where locally dominant salt molecule, not NaCl like in most evolved genotypes have higher relative oceans where oceanic conditions of approx. fitness in the local environment compared 30-40 PSU is presumably above their the other environment (Kawecki and Ebert, tolerance. As the species continues to 2004; Savolainen, Lascoux and Merilä, 2013), expand, areas hosting such conditions will is one likely reason for the success of an are likely to be met. One such area is the invading species. Local adaptations for ongoing establishment in Swedish waters. dispersal abilities have been shown to be First recorded in the country in 2008 especially frequent at the front of expanding (Karlskrona, 56°9′39″N 15°35′10″E), the ranges, where individuals experience less round goby has subsequently established intraspecific competition due to low itself in various parts of Sweden, including population densities. This has been shown in the archipelago of Karlshamn a classic example of an invasive species the (56°10′12″N 14°51′47″E) and Stockholm cane toad (Rhinella marina). Studies have (59°19′46″N 18°4′7″E), the islands Gotland shown changes in skeletal morphology, (57°30′N 18°33′E) and Öland behavior and even a reduced immune (56°44′N 16°40′E) as well as the harbor of system response, facilitating further Gothenburg (57°42′25″N 11°57′59″E) dispersal (Hudson et al., 2016; Brown, (Brockmark, 2016). Phillips and Shine, 2014; Brown and Shine, The Gothenburg estuary features a 2014). Front range specific changes also halocline where freshwater flows from the occur in the round goby as shown by Azour river Göta älv and saltwater pushes in from et al. (2015), with individuals at the forefront Kattegatt creating a complex and shifting of a distribution range exhibiting faster body of brackish water. Kattegat itself is growth and higher welfare due to less connected to the North Sea, which is an area intraspecific competition for food. Similarly, of higher