AREAS AND FORMING CENTERS OF THE GENERA Salmo, Oncorchynchus, Hucho AND Brachymystax (PISCES, SALMONIDAE) Simo Georgiev

ABSTRACT By comparing the areas of the genera Hucho, Brachymistax, Salmo and Oncorchynchus, as well as the geomorphology of Eurasia and North America and the positioning of young chain mountains around the North Pacific Ocean basin and South Europe, the author lays out a thesis on the reasons for forming of different number of species in the frame of those four genera in the family Salmonidae. From the mutual ancestors Hucho and Brachymistax that have appeared in Siberia, the former became a dominant predator which preferred the upper streams in the mountainous scenery while the latter became recessive, developed in smaller dimensions, feeding on invertebrates and preferring the still water or slow motion fluent habitats. The representatives of Salmo and Oncorchynchus have a mutual ancestor, similar biology and phenotypic appearance, and as a result, for a long time their species were placed together in the same genera. In both genera there are species which spend part of their life in salt waters in the North hemisphere. The reason for separation of their mutual ancestor in two genera, i.e. in West Eurasia Salmo and in East Eurasia and North America Oncorchynchus is the older freshwater Hucho that has conquered that area first. The extension of the Rocky Mountains in the South-North in parallel with the East seashore of the Pacific Ocean made it possible for the Oncorchynchus to form a larger number of species than in Salmo where the presence of the Alps, Pyrenees, Dinarids and Carpathians together with the unfavorable climate circumstances resulted with different development sequence. One only West-Eurasian Salmo species (S. salar) counter to the five North-American and East-Eurasian in Oncorchynchus (O. kisutch, O. tshawytscha O. keta, O. gorbuscha, O. nerka) exploit the rich forage of the world's ocean. The complicated geomorphologic orthography of South Europe in combination with the ice cover has caused different composition of salmonid habitats in the Mediterranean, especially on the Balkan Peninsula renowned for its endemic salmonids.] Key words: Salmoniformes, Salmo, Brachymistax, areal, forming center Oncorchynchus, Hucho,

2 INTRODUCTION The family Salmonidae has been subject to increasing interest for the long time mainly due to their economic importance (Vladykov, 1963). Some of the gens show big phenotypic plasticity as consequence of the poliploidy (Allendorf and Thorgaard, 1984, cited in Healey, 2009; Osinov and Bernachez, 1996). There is still no consensus on the exact number of species, or to be more precise, the status of some species (Rounsefell, 1962; Behnke, 1972; Smith and Stearley, 1989; Stearley and Smith, 1993; Kottelat and Freyhof, 2007); to some of the entities there are even approaches that are too emotional (Sunik et al. 2007). The rapid development of the molecular biology (Bernatchez et al. 1992; Osinov and Bernatchez, 1996; Apostolidis et al. 1997, 2011; Machordom et al. 2000; Bernatchez, 2001; Largiander et al. 2001; Froufe et al. 2003, 2004, 2005; Antunes et al. 2006; Campos et al. 2007; Martnez et al. 2007; Snoj et al. 2009; Weiss et al. 2011), made it possible to go into details when explaining the closest relationships among the separate populations. This debate is focused on the areals and forming centers of the freshwater-marine Salmo and Oncorchynchus as well freshwater Hucho and Brachymystax as indirect indicators for the events that have caused todays versatile number of species in the gens Oncorchynchus (Behnke, 1972) and Salmo (Bernatchez, 2001). The key conclusion for this debate resulted from the discover of Montgomery (2000) on the connection between the Pacific Ocean topography and evolutional radiation of Oncorchynchus; this discover was then reflected on the development destiny of Salmo, the species that have the largest areal Salmo trutta (Linnaeus, 1758). COMPARISON OF THE AREAS AND THE FORMING CENTERS The genera extension is shown on Fig. 1.

Fig. 1.

The debated genera areas extension

Sl. 1. Protegawe na arealite na razgleduvanite rodovi

The areas of Hucho (Vladykov, 1963; Holik, 1981, 1982; Weiss et al. 2011) and Brachymystax (Nikolsky, 1956; Berg, 1948; Vladykov, 1963; Holik, 1981) overlap in Asia (Siberia), while the first extends in Europe too but with some interruptions (in Danube), Fig. 2. There is a lack of convenient microhabitats in the larger surfaces at East of Danube and Volga's confluent Kama, in Dnieper, Dniester, Don and larger in the flow of Volga. The last listed rivers cross trough prairies and semi-deserts (Ozenda, 1994). It is also not present in the salmonid confluents of Danube in the Rhodope and

4 Haemus (Oresharov and Nishkov, 1959). The question of the absence in the habitats by convenient ecological characteristics in the mountainous confluents belonging to the watersheds of PontoCaspian depression, inhabited with Salmo, present together to Hucho in the flow of Danube (Drobnjakovi, 1934), still remains pending. The area of Hucho and Salmo in the middle Asia exclude each other in the watershed of Aral Sea. Although geologically old (Norden, 1961; Wilson, 1974; Kendall and Behnke, 1984; Dorofeeva, 1989, cited in Stearley and Smith, 1993, p. 3), only three species were developed in them. Even though they are members of same ichthyocenoses from zoogeographic point of view, ecologically they differ: Hucho is midely benthic fish eater hunting large prey and prefers the upper mountainous and sub mountainous parts of the rivers while Brachymystax is nektonic and pelagic mainly invertebrate eater and plankton eater; it demonstrates facultative fish predator features only in the oldest age classes, when they eat alevins or small prey (Mitrofanov and Petr, 1999). In other words, they are not competitors for food. When convenient circumstances appeared, they extended to the West, first the predecessor of Brachymystax (Berg, 1908; Hadzisce, 1961, cited in Stearley and Smith, 1993, p. 2) in the watershed of the Adriatic Sea, in Miocene (Hs, 1978, cited in Bianco, 1990, p. 172 "D Adriatic-Pannonian connection"), four million years ago (Sunik et al. 2006), later Hucho, in the flow of Danube (Georgiev, 2003). The areas of Salmo (Balon, 1968; Behnke, 1968; Bagliniere et al. 1994; Oakley and Phillips, 1999) and Oncorchynchus (Behnke, 1972), extend across both freshwater and saltwater habitats. Both species show plasticity, both phenotypic and ecologic; the plasticity of Salmo is especially expressed when found in artificial habitats, (Sidorovski, 1955, 1960, 1971). According to the results of Bernatchez (2001), Salmo has less species, the ambivalent Salmo salar Linnaeus, 1758 (Pedley and Jones, 1978; Nislow et al. 1998); the freshwater endemic Salmo obtusirostris (Heckel, 1851), Salmo ohridanus Steindachner, 1892, Salmo carpio Linnaeus, 1758, Salmo platycephalus Behnke, 1969 and Salmo marmorata Cuvier, 1829; the most expressed flexibility in feeding manner is manifested by Salmo trutta (Linnaeus, 1758), (Thomas, 1967; Debeljak and Faai, 1985; Nsje et al. 1998). The plasticity of S. trutta is expressed also during reproduction (Garca-Vazquez et al. 2001; Brabrand et al. 2002; Olsen and Vllestad, 2003). Most of the Oncorchynchus species are anadromous, nevertheless sometimes among the same species we come across exclusive freshwater populations (Behnke, 1972). This is the case of mid-large to large fish some of which are the largest in the order, and their vegetative phase in the Pacific is analogous to the one of S. salar in Atlantic. According to Stearley and Smith (1993), this is the genera closest to Salmo and the one that is most advanced in its evolution. It is richer in species (10), which results from the

5 matching features of the tectonic of the Pacific Ocean and the creation of the genus followed by accelerated evolutionary radiation (Montgomery, 2000). The North-East seashore of the Pacific, Alaska i.e. the Yukon river flow is considered as the center of forming of Oncorchynchus. The fossil Eosalmo one a "sister group of all living Salmonidae" (Wilson, 1977 cited in Stearley and Smith, 1993, p. 12) from Eocen, was found in the Rocky Mountains, South of the stream area of Yukon. From the North Eosalmo descended at South across the Pacific seashores of Asia and North America. Unlike Hucho that reached the Pacific by land, Oncorchynchus has reached to the Korean Peninsula, Japan Archipelago and Taiwan through water. This explains the richness in species spawning in the same place but does not give evidence of cruel competition. After the hatching, the alevins of two different species live together (Everest, 1972), one on the sandy part of the bottom and the others on the gravel bottom. Eventhough small in quantity, the primary production meets the needs of the earliest phases. Once the appropriate mass is reached, i.t. the mass that has needs exceeding the needs of the primary production, they go down in the more productive ocean. The center of forming of Salmo just S. trutta, was often subject to debates; two different positions are presented (Osinov and Bernatchez, 1996), according to one of these, this center is Scandinavia. I personally support the other thesis, which says that the center should be searched for in the south, the area of PontoCaspian depression. Unlike the authors cited in Osinov and Bernachez (1996), (Derzhavin, 1934; Vladimirov, 1944, 1948; Rukhkyan, 1989), instead of the Caucasus, I suggest Armenia, with very developed hydrographic network. Although S. trutta has a number of populations both in salt waters as far as in Iceland, which much nearer to the North America shores than to Armenian highland, nonetheless it prefers the small mountain brooks, representing the only ichthyofaunistic component. What leads us to this conclusion? The presence of Salmo in the Indian Ocean watershed, "behind" the high and large mountains chain separating the Ponto-Caspian depression and the Persian Gulf. It is easy to understand how a part of the S. trutta population from the Black Sea watershed, went up to the high waters of Taurus mountain, and hence in this seismic area, became a part of the Indian Ocean watershed. One additional contribution is the presence of Salmo deep in the middle Asia, Amu Darya and Sir Darya, recently cut off confluents of Aral Sea. S. trutta as the most typical representative of Salmo in the North West African seashore and recently in the West Mediterranean, "in at least three successive waves, 1.2, 0.4 and 0.20.1 MY ago" Snoj et al. (2011), in the Ohrid Lake only 155.000 of years ago (Sunik et al. 2007). As Scandinavia, the Alps and the area between them, during the ice period used to be glaciated so the living world was destroyed; some went down to the South in

6 some refugia, mainly on the Balkan Peninsula. Here we need to mention the "salinity crisis" (Bianco, 1990; Clauzon 2005) caused by connecting the most South-Western part of the European soil to the neighboring African land that stopped the immigration of the fish from the Atlantic to the Mediterranean Sea. The alternative thesis for Scandinavia as the center of Salmo formation would be the school that went across the already mentioned "D Adriatic-Pannonian connection" (Hs, 1978, cited in Bianco, 1990); from this school the S. marmorata Cuvier, 1829 was developed, it entered in the watershed of Adriatic/Mediterranean Sea; another school had colonized to the Ponto-Caspian basin and managed to reach the Indian Ocean watershed, when the third school conquered the fresh waters of Atlantic between Jiland and Pyrenees. The molecular researches do not support this thesis, they refer to the movements from East to the West across Middle Europe in some successive waves, and then through the Atlantic and Gibraltar from West to East in the Mediterranean (Sunik et al. 2007; Apostolidis et al. 2011). Consequently we can conclude that the populations by S. trutta in Scandinavia, Iceland and Mediterranean are younger than those in the Ponto-Caspian depression. From the mutual ancestor with S. marmorata which when the first wave crossed Scandinavia and the Alps at West and conquered the salty waters of the Atlantic Ocean, the S. salar developed; and a proof of that is the finding of the "ancestral state" of Apostolidis et al. (1997, p. 541) proving that there is the "the same nucleotid (A) . . . on the place 275". It is easy to explain the forming of S. salar in the North-East Atlantic. The slow incline of the soil rich in estuaries where the sweet water gradually becomes salted, has made it possible for the adaptation to occur of thee vegetative phase towards the more rich food in the marine water especially in the absence of competition (Hucho). That happened 4 MY ago, when the south school entered the Adriatic Sea together with the ancestor of S. obtusirostris and S. ohridanus (Bernatchez, 2001; Kottelat and Freyhof, 2007; Sunik et al. 2006, 2007, 2011; Snoj et al. 2008, 2009, 2011; Froufe et al. 2003, 2004, 2005), as Thymallus thymallus Linnaeus, 1758, (Zerunian, 2002), even though Osinov and Bernatchez (1996), locate the appearance of the ancestor of Salmo (S. trutta) some 2 MY ago. The reproduction in the sweet waters from Cape St. Vincent to Scandinavia is proof for the origin of S. salar. Part of S. trutta populations through the South confluents of Danube on the Balkan Peninsula, entered the neighboring flow of Vardar, Aegean Sea watershed, or at a larger scale, the North-East Mediterranean Sea, and from there in the other confluents of Aegean Sea, Mesta ("Nestos" in Bernatchez, 2001). S. trutta has different biological needs than Hucho which came from East last, so for it did not represent any danger as it was for S. marmorata with which it divides the same mountain mass on Balkan - Dinarids, and

7 the habitats on two different sides of the water separations. The absence of Salmo East of the Aral Sea, Ob, is due to the lack of the convenient ecological conditions in the recent time, as well as the presence of Hucho in the past. ANALOGIES AND DIFFERENCES AMONG GENERA Salmo AND Oncorchynchus The brilliant explanation offered by Montgomery (2000) on the way of forming such a large number of species in the genus Oncorchynchus, can help us understand why there are less species in the also so young genus Salmo. The axle of the Rocky Mountains chain extends in parallel to the East seashore of the Pacific Ocean distanced to 1000 km (+), creating a dense net of five large independent fluent habitats as well as a lot of small ones. During the ice periods, when the North habitats became unfavorable, the populations conquered new ones more to the South. For their vegetative phase they had on their disposal the largest water mass on the planet. In time some populations became independent species, going out from the ocean to spawn in different time or together but at different parts of the bottom. Therefore, there was no rivalry during reproduction and juvenile period, so separate species reproduced on the same place (Everest 1972). On the other hand, the extension of the mountains chain in Europe is different, so the development of the genus Salmo progressed in another manner. As we can see o Fig. 3., they lie on the East-West line or vice versa, so migrations/colonization after the closing of the Panonian-Adriatic connection could happen only in this line and only in the inter-glacial periods. This is the explanation why the species S. trutta express such large phenotypic differences. Having in mind the biology and the recent areas of S. trutta and S. salar as their dimensions, the scenery of their actual extension can be discovered very easily. From Armenia as proposed center of forming in accordance with Hillenius (1964), the ancestor with large dimensions could not extend on West because of the competition with the older and larger Hucho with whom they had the same biology, so the only free space for enlarging of the areal was to the West. The younger populations with smaller dimensions representing the recent complex S. trutta (Bernatchez, 2001; Mari et al. 2006, 2006a), migrated on West later, in waves during the inter-glacial periods, as explained above.

Fig. 2.

The exclusion, contact and overlap by the areas of the debated genera (excluding Brachymystax)

Sl. 2. Isklu~uvawe, dopirawe i preklopuvawe na arealite na razgleduvanite rodovi (so isklu~ok na Brachymystax)

A proof that Hucho caused separation of the mutual ancestor of Salmo and Oncorhynchus, as two genera of same ecology, is the strident competition among both the most extended and plastic species S. trutta and O. mykiss implanted in the same aquatoria by the anthropogenic factor (Brown et al. 1976; Gatz et al. 1987; Scott and Irvine, 2000). Today is difficult to say if besides Hucho, and the differences in the forage in the fluent habitats and in the wood and grass areas, had additional influence on the extension of the Salmo

9 and Oncorchynchus areas (Kawaguchi and Nakano, 2001). Comparing the discovery of Montgomery (2000) and the history of Salmo decoded by the newest molecular studies, it is easy to conclude that the different geomorphological conditions influenced differently on the evolution of genus Salmo from Oncorchynchus. On the African soil, on the south of the mountain Atlas the giant waterless space extends, the Sahara. The results of Sanz et al. (2000), demonstrate how the differences occur among the separate populations of S. trutta in the interglacial refugia.

Fig. 3.

The extension of mountain's chains in North America and South Europe

Sl. 3. Protegawe na planinskite sinxiri vo Severna Amerika i Ju`na Evropa

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14 Weiss, S., S. Mari, S., Snoj, A. (2011): Regional structure despite limited mtDNA sequence diversity found in the endangered Huchen, Hucho hucho (Linnaeus, 1758). Hydrobiologia, 658, 103-110. Zerunian S. (2002): Condannati all'estinzione Biodiversit, biologia, minacce e strategie di conservazione dei Pesci d'acqua dolce indigeni in Italia, Edagricole. + http://earth.google.com/geoeye/index.html ACKNOWLEDGMENT The crucial paper for this debate Vladykov (1963) kindly presented Anthi Oikonomou, Ioannina, doctorate student in ichthyology. Proofread by Mirjana Makedonska. REZIME Vrz osnova na sporeduvawe na arealite na rodovite Hucho, Brachymistax, Salmo i Oncorchynchus, kako i geomorfologijata na Evroazija i Severna Amerika, protegaweto na sinxirite na mladite planini okolu bazenot na severniot del na Pacifi~kiot Okean i ju`na Evropa, avtorot iznesuva teza za pri~inite na sozdavaweto na razli~en broj vidovi vo ovie ~etiri roda vo semejstvoto Salmonidae. Od zaedni~kiot predok na Hucho i Brachymistax pojaven vo Sibir, prviot stanal dominanten grablivec koj im dava prednost na gornite te~enija vo planinskite predeli, a vtoriot recesiven, so pomali dimenzii inverterbratofag, koj dava prednost na stoe~kite `iveali{ta i bavnote~e~kite `iveali{ta. Pretstavnicite na Salmo i Oncorchynchus imaat zaedni~ki predok, sli~na biologija i fenotipski izgled, {to pri~inilo nivnite vidovi dolgo vreme da bidat stavani vo ist rod. Vo obata roda ima vidovi koi del od `ivotot proveduvaat vo solena vodi na delovi od severnata polutopka. Pri~ina za podelba od nivniot zaedni~kipredok vo dva roda, vo zapadna Evroazija Salmo i vo isto~na Evroazija i severna Amerika Oncorchynchus e postariot slatkovoden Hucho so porano osvoeniot prostor. Protegaweto na Karpestite Planini vo nasoka sever jug naporedno so isto~noto krajbre`je na Tihiot Okean ovozmo`ilo Oncorchynchus da razvie pove}e vidovi otkolku Salmo kade protegaweto na Alpite, Pirineitea, Dinaridite i Karpatite vo sodejstvo so nepovolnite klimatski priliki predizvikalo poinakva nasoka na razvojot. Samo eden zapadnoEvroaziski vid vo Salmo (S. salar) nasproti pet severo-Amerikanski i isto~no-Aziskih kaj Oncorchynchus (O. kisutch, O. tshawytscha, O. keta, O. gorbuscha, O. nerka) ja polzuva bogatata krmna osnova na svetskiot okean. Komplikuvanata geomorfolo{ka ortografija na ju`ne Evropa zaedno so mrazovata pokrivka predizvikala poinakov sostav na pastrmskite `iveali{ta vo Sredozemjeto, osobeno na Balkanskiot Poluostrov poznat po endemi~nite pastrmki.

15 Klu~ni zborovi: Salmoniformes, Salmo, Oncorchynchus, Hucho, Brachymistax, areal, centar na sozdavawe