Trout and Char of Central and Southern Europe and Northern Africa
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12 Trout and Char of Central and Southern Europe and Northern Africa Javier Lobón-Cerviá, Manu Esteve, Patrick Berrebi, Antonino Duchi, Massimo Lorenzoni, Kyle A. Young Introduction !e area of central and southern Europe, the Mediterranean, and North Africa spans a wide range of climates from dry deserts to wet forests and temperate maritime to high alpine. !e geologic diversity, glacial history, and long human history of the region have interacted with broad climatic gradients to shape the historical and cur- rent phylogeography of the region’s native trout and char. !e current distributions and abundances of native species are determined in large part by their fundamental niches (i.e., clean, cold water with high dissolved oxygen). Brown Trout Salmo trutta are relatively common and widespread in the northern and mountainous areas of the region but occur in isolated headwater populations in the warmer southern areas of the region. !ese southern areas provided glacial refugia for salmonids and today har- bor much of the region’s phylogenetic diversity. Despite relatively narrow ecologi- cal requirements in terms of water quality, native and invasive trout and char occur throughout the region’s rivers, lakes, estuaries, and coastal waters. Despite having only a single widely recognized native trout species, the region’s range of environments has produced a remarkable diversity of life histories ranging from dwarf, stunted, short and long-lived, small- and large-sized, stream-resident, lake-resident, fluvial potamo- dromous, adfluvial potamodromous, and anadromous (see Chapter 7). Only one trout and one char are native to the region, Brown Trout and Alpine Char Salvelinus umbla. !ere are four currently recognized Brown Trout relatives from the Italian peninsula and Mediterranean islands—Salmo fibreni, S. carpio, S. cettii, and S. marmoratus—and a taxonomically complex and poorly studied group of Brown Trout relatives in Morocco and Algeria. !ree nonnative species from North America (sensu International Union for Conservation of Nature [IUCN]) have been introduced and established naturalized populations: Rainbow Trout On- corhynchus mykiss, Brook Trout Salvelinus fontinalis, and Lake Trout S. namaycush. !is chapter summarizes the information available on the distribution, status, and management of these trout and char. 1 2 12 Native Trout and Char Brown Trout Brown Trout is native throughout the region and is the most important species in terms of distribution, abundance, genetic and phenotypic diversity, and scientific, public, and economic interest. Brown Trout is one of the world’s most studied spe- cies and is used as a model system by population, behavioral, and evolutionary ecolo- gists and geneticists. Beyond the thousands of peer-reviewed journal articles, the vast literature on Brown Trout includes dozens of books, monographs, and symposia volumes. Newton (2013) and Lobón-Cerviá and Sanz (2017) provide contemporary summaries of Brown Trout biology and management. Phylogeography.—A dizzy array of colors and morphology typifies Brown Trout across its native range (see www.cartapiscicola.es or www.sibic.org/en/spanish-fish- chart/). Such variability in morphology, color, and life history has led to the nam- ing of more than 40 species! However, contemporary genetic data suggest that the single species Salmo trutta was modified by glaciation processes and colonization histories into multiple genetic lineages. Several lineages and sublineages in Italy, the Mediterranean islands, and North Africa have been elevated to species level. Fur- thermore, species designations are likely warranted for other sublineages in north- ern Europe (see Chapter 10), the Balkans, the Turkish peninsula, and northeastern Asia. Bernatchez et al. (1992) recognized five main genetic lineages for European Brown Trout: Atlantic trout (AT), Danubian trout (DA), Marble Trout of north Adriatic rivers (MA), Adriatic trout (AD), and Mediterranean trout (ME). Subse- quent studies (Bernatchez 2001; Cortey and García-Marín 2002) highlighted sev- eral genetically distinct sublineages with some of these main lineages. !ese include the Duero lineage (DU) from the Iberian Peninsula (Suarez et al. 2001), the Dades lineage (DAD) in North Africa (Snoj et al. 2011), and the Tigris sublineage (TI; Bardakci et al. 2006) from the Euphrates River (Turkey). Two sublineages, Salmo obtusirostris and S. ohridanus, have been recently reclassified into the genus Salmo as distinct species (Phillips et al. 2000; Sušnik et al. 2006; Snoj et al. 2009). Six of these lineages occur in the region covered by this chapter: 1. !e AT lineage occurs in rivers from Norway to the Iberian Peninsula and Sicily (Schöffmann et al. 2007; Cortey et al. 2009; Fruciano et al. 2014). Several mor- phological species have been described within the AT lineage (gillaroo Salmo sto- machicus Günther 1866; sonaghen S. nigripinnis Günther 1866, and S. ferox Jar- dine 1835). Sicilian trout were described as S. cettii Rafinesque 1810 but later on assigned to the AT lineage (Schöffmann et al. 2007; Fruciano et al. 2014). !e AT lineage has been widely stocked outside its native range but is considered native to the upper Danube (Schenekar et al. 2014) and is probably native in some north Italian rivers draining to the Mediterranean (Meraner et al. 2007). / 3 2. !e DU lineage is native to the Duero and Miño rivers in the Iberian Peninsula (Vera et al. 2010). 3. !e ME lineage has a native range in the Mediterranean rivers of the Iberian Peninsula (Cortey et al. 2004), southern France and Corsica (Berrebi et al. 2000; Berrebi 2015), and Italy and Greece (Giuffra et al. 1994; Bernatchez 2001). 4. !e MA lineage is native to rivers of the northern Adriatic basin (Bernatchez et al. 1992; Giuffra et al. 1994; Snoj et al. 2000). !e recent revision by Pustovrh et al. (2014) assigns the species to this MA lineage. 5. !e AD lineage is distributed along the Mediterranean rivers from the southern Iberian Peninsula to Turkey, including Corsica (Cortey et al. 2004; Snoj et al. 2011; Berrebi 2015). Several morphological species endemic to the Italian penin- sula (Salmo carpio Linnaeus 1758; S. cenerinus Kottelat 1997, S. cetti Rafinesque 1810, and S. fibreni Zerunian & Gandolfi 1989 [Gratton et al. 2014]) and the Balkans and Turkish Peninsula (S. dentex Heckel 1851, Flathead Trout S. platyc- ephalus Behnke 1969, S. macrostigma Duméril 1858, S. lourocensis Delling 2011, Ohrid Trout S. letnica Karaman 1924, S. peristericus Karaman 1938, and S. pela- gonicus Karaman 1938) are members of the AD lineage (Sušnik et al. 2004, 2007; Lo Brutto et al. 2010; Snoj et al. 2011). 6. !e DA lineage is native to the Danube basin and Vistula River flowing to the At- lantic (Kohout et al. 2012). Several morphological species (Sevan Trout Salmo ischchan Kessler 1877, Tigris Trout S. tigridis Turan et al. 2011, Black Sea Salmon S. labrax Pallas 1814, Caspian Salmon S. trutta caspius Kessler 1887, and Amu- Darya Trout S. t. oxianus Kessler 1874) are members of the DA lineage. Within and among these lineages, Brown Trout exhibit several life-history forms. !ese forms, generally known as resident, anadromous, and lacustrine-adfluvial, have recently being reviewed according to Varley and Gresswell (1988) and Northcote (1997) as follows. Fluvial populations are where reproductive, feeding, and refuge mi- grations occur in the rivers and streams of the home range. Fluvial-adfluvial popu- lations migrate from main stems into tributaries to spawn, but progeny may move downstream for growth and refuge. Lake-dwelling populations may spawn in inlets (lacustrine-adfluvial migration pattern), outlets (allacustrine migration pattern), and the lake itself (nonmigratory lacustrine spawners). Nevertheless, life-history expressions is a complex topic of much ongoing inves- tigation because it is a heritable but plastic trait and many fish express alternative life histories in partial sympatry and interbreed. For example, within a river draining into the Atlantic Ocean, the ratio of anadromous to resident trout typically decreases with distance from the sea and anadromy is more common among females assumedly be- cause fitness of females depends more on body size than males. Similarly, in a single river, resident forms may predominate in headwaters and adfluvials in the main-stem rivers draining into lower elevation lakes. We recognize this life-history diversity and distinguish among forms throughout the text. 4 12 Distribution.—!e naturally complex distribution of Brown Trout lineages and life-history forms in central and southern Europe has been modified through habitat fragmentation and stocking of nonnative lineages. For example, in the Czech Repub- lic, more than 500 km from the nearest estuary, anadromous forms occurred until the damming of the Elbe and Oder rivers in the early 20th century, but now only resident trout occur in the headwaters of the Elbe, Danube, and Oder (Kohout et al. 2012). Although there is evidence that the distributions of lineages naturally overlap in some basins (Bernatchez 2001; Lerceteau-Köhler et al. 2013), the range of the AT lineage, in particular, has expanded dramatically through stocking. In Germany, resident trout are common in high altitude streams, adfluvial individ- uals occur in subalpine lakes and large reservoirs, and anadromous forms predominate in the North Sea drainages of Schleswig-Holstein, Lower Saxony, Saxony-Anhalt and North Rhine-Westphalia. Historically, anadromous forms were present up to the Bohemian tributaries of the Elbe and Upper Rhine. Six rivers currently have sea trout: the Eider, Elbe, Weser, Ems, Meuse, and Rhine. !e Kiel Canal, built in 1890, allowed historically isolated anadromous populations from the Baltic and North Sea basins to hybridize (Petereit et al. 2015). In neighboring Netherlands, anadromous trout were likely present historically, but now, only resident forms remain in coldwater refugia of severely degraded and fragmented channel networks. In Switzerland, the presence of five genetic lineages, complex topography, and long history of stocking exemplify how natural and anthropogenic factors interact to affect the distribution of Brown Trout lineages and life-history forms.