Morphological Differentiation of Salmonines (Subfamily Salmoninae) with Emphasis on Trout Salmo Spp

See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/259936344 Morphological Differentiation of Salmonines (Subfamily Salmoninae) with Emphasis on Trout Salmo spp. Stocks in Serbia and Adjacent Regions Article in Acta Zoologica Bulgarica · January 2005 CITATION READS 1 136 3 authors, including: Predrag Simonović Saša Marić University of Belgrade University of Belgrade 159 PUBLICATIONS 1,598 CITATIONS 95 PUBLICATIONS 711 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: “Climate change and invasive species- assessing effects onto biodiversity of native freshwater crayfish and salmonids and their conservation” View project HydroMediT 2021 View project All content following this page was uploaded by Predrag Simonović on 01 February 2014. The user has requested enhancement of the downloaded file. ACTA ZOOLOGICA BULGARICA (KARAMAN, 1927), occurring in sympatry in different areas of the lake basin. In addi- Acta zool. bulg., 57 (3), 2005: 341-362 tion, the distinct West-Balkan endemic genus Salmothymus BERG, 1908 (including Acanthiolingua HADIÈE, 1960), with several stocks in Lake Ohrid and rivers in the Adriatic Sea Basin contributes to the diversity of salmonine fish fauna of the region. The variability of external morphological characters in trouts was extensively stud- ied in the Balkans (HECKEL, KNER 1858, KARAMAN 1926, 1932, 1934, 1938, KARAKOUSIS Morphological Differentiation of Salmonines et al. 1991, DELLING 2002, 2003a, 2003b). However, the majority of references are so (Subfamily Salmoninae) with Emphasis on Trout old and the analytical approach they contain out-of-date that they are not in citing any more. KARAKOUSIS et al. (1991) failed to distinguish brown trout stocks of Greece for Salmo spp. Stocks in Serbia and Adjacent Regions their external morphology, whereas DELLING (2002, 2003a) revealed the distinctness of marble trout and several other brown trout taxa from the Balkans in their morphologi- Predrag Simonoviæ1, Saa Mariæ1 and Vera Nikoliæ1 cal characters and analyzed their phylogenetic status. The rapid progress in biochemical and molecular methods enabled the clear definition of particular phyletic distinct lin- Abstract: The multivariate analysis of the external morphology of 22 continuous characters eages of the brown trout, e.g., Atlantic, Danubian, Mediterranean, Adriatic and reveals that huchen Hucho hucho and Arctic charr Salvelinus alpinus are different from all marmoratus (BERNATCHEZ et al. 1992, BERNATCHEZ 2001), including the revelation of trout stocks, whereas the Lake Ohrid belvica Salmothymus ohridanus is close to the brown their evolutionary history. In such recognized main lineages, however, particular isolates trout complex. The position of rainbow trout Oncorhynchus mykiss implies greater similarity remained still insufficiently explained in a sense of taxonomy (KOTTELAT 1997), regard- in trout-salmon morphology in relation to that of huchen and charr. The phenetic relation- ing the discrepancy between the molecular data and those coming from the investigation ships derived on external morphology suggest similarity of the west-Danubian brown trout on morphology. Particular opinions, e.g. that of K (1997), have reservations stock of the Black Sea basin to the brown and marble trout stocks from the Adriatic Sea OTTELAT basin, difference of brown trout stock from the Mlava River and similarity of brown trout about the initially given significance of the results in the field of taxonomy based stock from the Velika Morava River drainage in comparison to that of the Aegean Sea exclusively on biochemical and molecular data, without taking into account the data basin. Those results are initial hints for the conservation of brown trout stocks of Serbia. coming from the analyses of morphology, zoogeography and distribution. Moreover, DELLING (2002) stated that molecular markers widely applied in salmonid biology have a Key Words: phenetics, evolutionary history, river drainages, brown trout stocks potential to falsify recognizable species. This criticism came out from the much scarcer contribution to the systematics of fish from the molecular and biochemical analyzes than Introduction expected. Regardless of how much biochemical and molecular analyses attempt to replace the comprehensive analysis of morphophenotype with the analysis of particular minor The problem in brown trout taxonomy comes both from the great phylogenetic oldness features of genome, it seems that phenetics associated with sophisticated analytic tools can of brown trout and from the dynamic evolutionary history in various parts of broad still give its contribution in clarification of trout stocks taxonomy. dispersal area. The Balkan Peninsula is known for its nominal brown trout diversity. The initial step for getting an insight into the diversity of brown trout stock of However, the taxonomic status of those trouts is still unclear. KARAMAN (1926) stated particular river drainages and sea basins of the Balkans is the investigation of the that the isolation of brown trout populations dating from the Tertiary (e.g., Lake Ohrid variability of their external morphology by exploratory techniques that should indicate trout taxa) is a consequence of mountain chains genesis (e.g., Dinarid Alps), whereas the basic level of diversity and facilitate the decision-making about further exploratory the ice ages (within-drainage brown trout taxa) isolation after Pleistocene is the result of steps. Considering that stocks of the Balkan brown trout are under the utilization of the refuge character of the Balkans during and after the Quarternary ice-ages. The recreative fisheries and that they are already long-term isolated both within the same or dynamic geological history of the area is reflected in prominent taxonomic diversity, between different river drainages of Serbia (e.g., those of Drina, Morava, Mlava, Timok together with the rather uniform ecological pattern of brown (including the marble) rivers and some other, smaller rivers flowing into the Sava and the Danube rivers), there trout stocks throughout the Balkan Peninsula. Brown trout stocks in the West Balkans is a strong conservational need to get an insight into their original diversity before it is are the Black Sea basin stationary brown trout taxa Salmo trutta L., 1758, and / or S. severely disturbed by fish stocking. taleri (KARAMAN 1932), Aegean Sea basin stationary brown trouts S. macedonicus KARAMAN, 1924, and S. pelagonicus KARAMAN, 1938, Adriatic Sea basin stationary Materials and Methods brown trout taxa S. farioides KARAMAN, 1938, S. dentex HECKEL, 1852, and marble trout S. marmoratus CUVIER, 1829, as well as several Lake Prespa and Ohrid brown Majority of samples of brown trout were collected from the West Balkans, i.e., from the trout taxa, e.g., S. peristericus KARAMAN, 1938, Salmo letnica (KARAMAN, 1924), Salmo streams situated in East and West Serbia (cf. S. trutta), Bosnia and Herzegovina, Montenegro lumi P OLJAKOV; FILIPI, BASHO, 1958, Salmo aphelios K OTTELAT, 1997, and S. balcanicus and FYR Macedonia (Fig. 1). These were obtained using electrofishing gears (samples S2, S3, S5, S6, S8 and SM) and by angling (samples S1, S4 and SZ) in the period from May 1996 to October 1999. The rest of brown, marble and rainbow trout samples were 1Faculty of Biology, University of Belgrade, Studentski trg 16, 11000 Belgrade, Serbia and Montenegro 341 342 Table 1. Samples of salmonid fish, with their locality names, taxonomical assignments, sample sizes (N), geographical determination of localities from which samples originate, notification on particular specifici- ties and abbreviations used in the paper. Area, drainage, sea Locality Taxon N Note Abbr. basin Herzegovina, Neretva Trešnjica fish Buna River Salmo farioides 12 S7 River, Adriatic Sea farm Trebuèica Salmo Slovenia, Soæa River, 18 Tolmin fish farm SB River marmoratus Adriatic Sea Salmothymus FYR Macedonia, Crni Lake Ohrid 6 SO ohridanus Drim River, Adriatic Sea Western Serbia, Sava Trešnjica fish Drina River Hucho hucho 11 HH River, Black Sea farm Oncorhynchus Eastern Serbia, Danube agubica fish Mlava River 14 OM mykkis River, Black Sea farm Salvelinus Bosnia, Sava River, Introduced - Lake Boèac 10 SA alpinus Black Sea stocked Montenegro, Skadar Upstream of Zeta River Salmo taleri 10 SZ Lake, Adriatic Sea subterranean pass Salmo Southern Serbia, Struma Boica River 25 SM macedonicus River, Aegean Sea Eastern Serbia, Velika Resava River Salmo trutta 9 S6 Morava River, Black Sea Godljevaèa Western Serbia, West Salmo trutta 13 S1 River Morava River, Black Sea Eastern Serbia, Mlava Upstream of Buk Stream Salmo trutta 17 S2 River, Black Sea subterranean pass Eastern Serbia, Danube Mlava River Salmo trutta 9 S3 Fig. 1. Sampling area and localities in the West Balkans. River, Black Sea Krupaja Eastern Serbia, Mlava Salmo trutta 5 S4 River River, Black Sea provided from fish farms (Table 1). In addition to the brown trout, samples of several Central Serbia, West Bela River Salmo trutta 7 S5 salmonid taxa, e.g., Lake Ohrid belvica Salmothymus ohridanus (STEINDACHNER, 1892), Morava River, Black Sea rainbow trout Oncorhynchus mykkis (WALBAUM, 1792), huchen Hucho hucho (L., 1758) Western Serbia, Drina Crni Stream Salmo taleri 10 S8 and charr Salvelinus alpinus (L., 1758) were included into the analysis as outgroups. The River, Black Sea samples comprised 176 individuals totally, the majority of them was sexually

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