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New Endemic Deepwater Charr Morphs of the Genus Salvelinus (Salmoniformes: Salmonidae) from Lake Kronotskoe, Kamchatka G

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New Endemic Deepwater Charr Morphs of the Genus Salvelinus (Salmoniformes: Salmonidae) from Lake Kronotskoe, Kamchatka G ISSN 1063-0740, Russian Journal of Marine Biology, 2017, Vol. 43, No. 3, pp. 216–223. © Pleiades Publishing, Ltd., 2017. Original Russian Text © G.N. Markevich, E.V. Esin, E.A. Saltykova, O.Yu. Busarova, L.A. Anisimova, K.V. Kuzishchin, 2017, published in Biologiya Morya. ORIGINAL PAPERS Ichthyology New Endemic Deepwater Charr Morphs of the Genus Salvelinus (Salmoniformes: Salmonidae) from Lake Kronotskoe, Kamchatka G. N. Markevicha, *, E. V. Esina, b, E. A. Saltykovac, O. Yu. Busarovad, L. A. Anisimovab, and K. V. Kuzishchine aKronotsky State Natural Biosphere Reserve, Yelizovo, 684000 Russia bAll-Russian Research Institute of Fisheries and Oceanography, Moscow, 107140 Russia cSevertsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, 119071 Russia dFar Eastern State Technical Fisheries University, Vladivostok, 690087 Russia eDepartment of Biology, Moscow State University, Moscow, 119234 Russia *e-mail: [email protected] Received December 1, 2016 Abstract⎯Two previously unknown profundal dwelling charr morphs of the genus Salvelinus from Lake Kro- notskoe are described in this article. According to their lifestyle peculiarities, these morphs were named the “bigmouth charr” and “smallmouth charr.” The former group is near-bottom benthivorous, while the latter one inhabits the water column and occupies the omnivorous niche. Bigmouth and smallmouth charrs are dis- tinguished from the rest of the sympatric charr morphs by their smaller size, shorter snout, larger eyes, as well as by several craniological traits and lower parasite abundance. Spawning of both morphs takes place directly in the profundal zone and lasts from late October to February. Additional studies are required to determine the systematic status of the new morphs. Keywords: microevolution, sympatric speciation, trophic-based specialization, profundal zone DOI: 10.1134/S1063074017030087 INTRODUCTION niches of the littoral zone and epilimnion were previ- Charrs of the genus Salvelinus are an extremely ously described in this water body [3, 11]. Subse- flexible group of salmonids. Their diversification is quently, a dwarf charr was described from the source of determined by resource utilization in distinct food the Kronotskaya River [8, 22]. It was also proposed to niches, which is connected with the evolution of spe- consider Dolly Varden from the lower reaches of the cific morphological traits and results in the origin of lake tributaries as an independent morph [7, 22]. similar morphs in numerous lacustrine–riverine sys- No catches were conducted previously in the pro- tems of the Holarctic [17]. The most widespread diver- sification pathway is specialization along the pelagic– fundal zone of Lake Kronotskoe for various reasons. benthic resource axes [15], which basically results in This lake is one of the deepest water bodies in the the benthivorous and planktivorous, or predatory, region (with its maximum depth of 136 m and the morphs [19]. Apart from relatively simple examples of average depth of 58 m). The majority of sediments in intralacustrine diversification, the genus Salvelinus the profundal zone of the lake basin are composed of charrs exhibit from three to five morphs in rare cases silt reaching a thickness up to 12 m, while the extensive [20, 23]. In particular, in addition to the shallow-water coastal slopes are composed of unsilted lava boulders. morphs, large and deep lakes are also inhabited by During the profundal zone surveys in 2012–2014, it charrs in the profundal zone. Such specific morphs was originally established that the deepwater part of have been described for S. namaycush (Walbaum, the lake is inhabited by two previously unknown charr 1792) from the Great Lakes of North America [18, 24], morphs. These new morphs were named the “big- for S. alpinus (Linnaeus, 1758) from lakes of Scandina- mouth” and “smallmouth” charrs. The skull mor- via [19], Transbaikalia [16], and the Taimyr Peninsula phology, ecology, and parasite fauna of these morphs [6]. A high variability in trophic and ecological char- were briefly described as a result of the first catches [1, acteristics was found for charrs in Lake Kronotskoe on 5, 12–14]. The objective of this study is to provide a the Kamchatka Peninsula. Three charr morphs that detailed description of the morphology, feeding pecu- occupy the benthivorous (nosed charr), predatory liarities, and lifestyle of the recently discovered pro- (longhead charr), and omnivorous (white charr) fundal charr morphs in Lake Kronotskoe. 216 NEW ENDEMIC DEEPWATER CHARR MORPHS OF THE GENUS SALVELINUS 217 Table 1. The morphometric characteristics (mean ± error of on lines of an estimated length were applied to catch the mean/lim) of deepwater charrs of the genus Salvelinus fish in the water column. A total of 516 bigmouth and from Lake Kronotskoe 403 smallmouth charrs were sampled. The length and Traits Bigmouth charr Smallmouth charr weight of all individuals were measured and the sex was determined. In 30 fish of each morph the age was FL*, mm established using polished otoliths (the opaque zone 274± 1.7 204± 1.4 edge was considered as annual ring). In 50 fish of each 240–358 168–330 morph, morphometric measurements were conducted by the standard scheme [9]. Variations in skull mor- FL, % phology (chondrocranium and 11 bones) were quanti- ± ± fied in 20 fish of each morph. The cranial traits were c* 19.7 0.17 18.9 0.17 18–23 17–21 evaluated based on the principle that was applied ear- ± ± lier to the epilimnic morphs of Lake Kronotskoe aD 42.1 0.14 41.7 0.16 charrs [3]. In 30 fish of each morph, the food compo- 40–45 39–45 sition was determined as the proportions in stomach of ± ± pD 40.5 0.16 41.4 0.19 different prey groups [10]. In the same fish, parasite 38–43 39–44 species composition and abundance [1], as well as ± ± infection intensity, were assessed by complete parasi- aV 48.5 0.18 48.4 0.22 46–52 45–51 tological dissection [2]. 67.4± 0.13 67.6± 0.14 To assess the statistical significance of morpho- aA metric and size–weight differences between the fish 66–68 66–69 groups, the Student’s t-test was used after checking the ± ± P–V* 29.4 0.20 30.5 0.25 normality of the characteristics by the Kolmogorov– 27–32 28–34 Smirnov test. The differences in diet and parasite ± ± infection were validated by the Mann–Whitney U-test V–A 20.5 0.16 20.5 0.18 19–23 18–23 [4]. Mathematical processing of the data was carried out in the Statsoft Statistica 13.0 software package. ± ± lP* 16.6 0.16 14.5 0.11 15–18 12–17 ± ± RESULTS AND DISCUSSION lV* 12.0 0.12 10.6 0.09 10–14 8–12 External View B“c”, % The bigmouth charr is characterized by a massive ± ± ao* 17.0 0.34 15.7 0.29 head with protruding supraorbital arcs; eyes are shifted 13–22 12–20 towards the nape, their average diameter is 27% of 26.8± 0.33 31.5± 0.30 head length. Upper jaw extends far behind the poste- o* rior eye edge, reaching 50% of head length on average; 22–31 29–35 lower jaw is long, curved, protruding forward and ± ± lmx* 49.9 0.55 44.5 0.43 reaching 70% of head length on average that provides 44–57 40–50 the upper mouth position. Body is cylindrical; caudal ± ± peduncle is long and low. Pectoral and ventral fins are lmd* 69.9 0.51 62.7 0.63 62–79 54–70 long and wide; pectoral fins length is greater than 0.5 of the P–V distance (Table 1). Caudal fin is slightly ± ± cH* 70.5 0.71 73.3 0.75 forked or truncated (Fig. 1a). Meristic characters are 50–79 65–84 as follows: D 8–10 (9), A 7–9 (8), P 12–14 (13), V 8– * The differences between the morphs are significant (t-test, p < 9 (8) branched rays; ll 123–132 (127); pc 27–42 (33); 0.05). rb 10–12 (11); sp. br. 15–18 (16); vert 61–68. FL, body length; c, head length. Distances are as follows: aD and pD, ante- and postdorsal; aV, anteventral; aA, anteanal; P–V, pecto- Mature individuals are characterized by dark flanks ventral; V–A, ventro-anal; lP and lV, length of pectoral and ven- and a reddish belly; parr spots are absent. Spots tral fins. Head measurements are as follows: ao, snout length; o, (always smaller than the pupil diameter) below the lat- eye diameter; lmx and lmd, length of upper and lower jaws; eral line are red or pink; above the lateral line, pink or cH,head height. white; no spots are found on fins. Jaws are usually uncolored, sometimes with red bordering along the MATERIALS AND METHODS edge. Gill rays have a deep black pigmentation. All fins are red; the first ray of pectoral and ventral fins, as well Catches were carried out from May to November as the lower rays of caudal fin are white. Bigmouth 2012–2014 in the profundal, limnetic, and littoral charr acquire the characteristic traits at an early age zones of the lake using a set of gill nets with a mesh size and are well identified by the curved lower jaw begin- from 18 to 35 mm. Floating gill nets with weights fixed ning from a body length (FL) of 7–10 cm. RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 43 No. 3 2017 218 MARKEVICH et al. (a) (b) Fig. 1. The external form of bigmouth (a) and smallmouth (b) charrs from the profundal zone of Lake Kronotskoe. Scale bar, 5 cm. The smallmouth charr head is mostly conic, wedge-shaped protrusion anteriorly to ventral fins.
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