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Carassius Ve Střední Evropě

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Carassius Ve Střední Evropě MASARYKOVA UNIVERZITA Přírodovědecká fakulta Ivo PAPOUŠEK MOLEKULÁRNĚ-GENETICKÉ ANALÝZY DRUHŮ RODU CARASSIUS VE STŘEDNÍ EVROPĚ Přílohy k disertační práci Školitel: prof. RNDr. Jiřina Relichová, CSc. Školitel-specialista: RNDr. Věra Lusková, CSc. Brno, 2008 PŘÍLOHY – SOUBOR PUBLIKACÍ Příloha I Papoušek, I., Vetešník, L., Halačka, K., Lusková, V., Humpl, M. & Mendel, J. (2008). Identification of natural hybrids of gibel carp Carassius auratus gibelio (Bloch) and crucian carp Carassius carassius (L.) from lower Dyje River floodplain (Czech Republic). Journal of Fish Biology 72, 1230-1235. Příloha II Vetešník, L., Papoušek, I., Halačka, K., Lusková, V. & Mendel, J. (2007). Morphometric and genetic analysis of Carassius auratus complex from an artificial wetland in Morava River floodplain, Czech Republic. Fisheries Science 73, 817-822. Příloha III Papoušek, I., Lusková, V., Halačka, K., Koščo, J., Vetešník, L., Lusk, S. & Mendel, J. (2008). Genetic diversity of the Carassius auratus complex (Teleostei: Cyprinidae) in the central Europe. Journal of Fish Biology (submitted 10/2008). Příloha I Papoušek, I., Vetešník, L., Halačka, K., Lusková, V., Humpl, M. & Mendel, J. (2008). Identification of natural hybrids of gibel carp Carassius auratus gibelio (Bloch) and crucian carp Carassius carassius (L.) from lower Dyje River floodplain (Czech Republic). Journal of Fish Biology 72, 1230-1235. Journal of Fish Biology (2008) 72, 1230–1235 doi:10.1111/j.1095-8649.2007.01783.x, available online at http://www.blackwell-synergy.com Identification of natural hybrids of gibel carp Carassius auratus gibelio (Bloch) and crucian carp Carassius carassius (L.) from lower Dyje River floodplain (Czech Republic) I. PAPOUSˇEK*, L. VETESˇNI´K,K.HALACˇKA,V.LUSKOVA´,M.HUMPL AND J. MENDEL Department of Ichthyology, Institute of Vertebrate Biology, Czech Academy of Sciences, v.v.i., Kveˇtna´ 8, 603 65 Brno, Czech Republic (Received 13 July 2007, Accepted 28 November 2007) Morphometric and genetic variation were examined in parental Carassius auratus gibelio and Carassius carassius individuals, and their natural hybrids. Of meristic traits, only the number of gill rakers clearly distinguished hybrids (39Á4 Æ 1Á4) from parental C. a. gibelio (48Á3 Æ 0Á7) and C. carassius (28Á6 Æ 1Á1). MtDNA sequences showed that the hybrids were descendants of female C. a. gibelio. Microsatellite analysis confirmed the presence in hybrids of variants typical of C. a. gibelio and C. carassius. # 2008 The Authors Journal compilation # 2008 The Fisheries Society of the British Isles Key words: crucian carp; gibel carp; gill rakers; hybrids; microsatellites; mtDNA. The genus Carassius is represented in Europe by four taxa, native crucian carp Carassius carassius (L.), introduced goldfish Carassius auratus auratus (L.), gibel carp Carassius auratus gibelio (Bloch) and rare ‘ginbuna’ Carassius auratus langsdorfii Temminck & Schlegel (Szczerbowski, 2002; Kalous et al., 2007; Vetesˇnı´k et al., 2007). Even though C. a. gibelio and C. carassius are widely dis- tributed, published reports of hybrids between them are rare. Kux (1982) described the occurrence of such hybrids from the Danube Delta, and Goryunova & Skakun (2002) found hybrids of these taxa in natural lakes in Kazakhstan. In Polesi waters (Ukraine), several hybrids were found by Mezherin & Liseckij (2004). Ha¨nfling & Harley (2003) and Ha¨nfling et al. (2005) report the frequent occurrence of diploid hybrids arising from C. auratus and C. carassius in Great Britain. Despite the general prevalence of C. a. auratus and C. a. gibelio in Europe, information on the occurrence of natural hybrids with native C. carassius is scarce. This limited information may not reflect the *Author to whom correspondence should be addressed. Tel.: þ420 543 422 529; email: [email protected] 1230 # 2008 The Authors Journal compilation # 2008 The Fisheries Society of the British Isles IDENTIFICATION OF NATURAL CARASSIUS HYBRIDS 1231 extent of hybridization or may indicate that insufficient attention has been paid to hybrid identification. The aim of the present study was to identify with mor- phometric and genetic analyses natural hybrids of gibel carp and crucian carp in the lower Dyje River floodplain (Czech Republic). A total of 634 individuals attributed to genus Carassius [visually identified as C. a. gibelio (n ¼ 594), C. carassius (n ¼ 25) and possible hybrids (n ¼ 15)] were caught by electrofishing in the parapotamon side arm in the aluvium of the lower Dyje River (48°439250 N, 16°539200 E; in an area of 0Á45 ha). Analyses of ploidy levels were performed by means of computer-assisted microscopic image cytometry according to Flajsˇhans (1997) and Halacˇka & Luskova´ (2000). Blood smears were stained by Harris hematoxyline (Sigma-Aldrich, Prague, Czech Republic) for 10 min. Ploidy diagnostic dimension (i.e. erythrocyte nuclear area) was determined by bright field microscopy, imaging by a charge-coupled device (CCD) camera and image processing in Olympus MicroImage 4.0 analyser. The ploidy status was determined in 75 fish, including C. a. gibelio [17 diploid males (m), 15 diploid and 43 triploid females (f)], 10 C. carassius (6 m and 4 f) and 15 hybrids (6 m and 9 f). Meristic traits were assessed according to Holcˇı´k (1989) in 12 individuals (6 m and 6 f) of diploid C. a. gibelio, eight individuals (4 m and 4 f) of C. carassius and 12 hybrids (5 m and 7 f). For the meristic analyses, individuals were killed by overdose of anaesthetic (2-phenoxy-ethanol) and preserved in 4% formaldehyde. Differences in values of meristic traits among C. a. gibelio, C. carassius and hybrids were tested using ANOVA in Statistica Cz 7 and detrended correspondence analysis (DCA) Canoco 3.10. All data were checked for normalcy before analyses. DNA analyses were performed on two C. carassius individuals (1 m and 1 f), two C. a. gibelio individuals (1 m and 1 f) and nine putative hybrids (4 m and 5 f). Tested fish were fin-clipped and the clips were stored in ethanol for genetic analyses. Total genomic DNA was isolated from fin clips by phenol-chloroform extraction according to Sambrook et al. (1989) with minor modifications. Frag- ments of mtDNA representing the hypervariable part of the control region (D-loop) were amplified using primers CarU32 (59-CCAAAGCCAGAATTCTAAAC-39) and CarL509 (59-CATGTGGGGTAATGA-39). Polymerase chain reaction (PCR) was performed in total volume of 50 ml, which contained 200 mMofeach primer, 200 mM dNTP, 1Á5 mM MgCl2,1Â reaction buffer, 100 ng of total genomic DNA and 1Á5 U Taq polymerase. The following protocol for amplifica- tion was used: denaturation 95° C 1 min, 32 cycles 94° C 45 s/50° C30s/72° C 45 s, final extension 72° C for 7 min. PCR fragments were purified using poly- ethyleneglycol (PEG)/MgCl2/NaAc and sequenced on an automatic sequencer ABI PRISM 310 (Applied Biosystems, Foster City, CA, U.S.A.). Sequences (493 bp) were aligned by MEGA 3.1 (Kumar et al., 2004) and compared with GenBank sequences of C. a. gibelio (accession number AJ 388413.1), C. a. auratus (NC 006580), C. a. langsdorfii (AB 052327.1) and Carassius auratus cuvieri Temminck & Schlegel (AB 052334.1), and unpublished sequences of C. carassius. Microsatellite loci GF17, GF29, MFW2 and MFW7 were amplified with previ- ously published primer sequences (Zheng et al., 1995; Crooijmans et al., 1997). PCR was performed according to Ha¨nfling et al. (2005). Resulting genotypes were compared with allele size ranges of C. carassius and C. a. gibelio as in Ha¨nfling et al. (2005). # 2008 The Authors Journal compilation # 2008 The Fisheries Society of the British Isles, Journal of Fish Biology 2008, 72, 1230–1235 1232 I. PAPOUSˇEK ET AL. Carassius carassius body colour was greenish with a golden tinged, luteous abdomen; C. a. gibelio had a dark grey back and silvery sides. Body colouring of hybrid individuals was similar to C. a. gibelio, with a tinge of dark grey and black on head, torso and fins. The upper line of the dorsal fin was convex and the upper edge was rounded in C. carassius, whereas in C. a. gibelio the upper line of the dorsal fin was ventrally slightly concave. The dorsal fin of hybrid individuals merged traits of both parental species: the cranial half of the dorsal fin was convex and corresponded to the fin of C. carassius; the caudal half of the dorsal fin was slightly concave as in C. a. gibelio. The inner side of the abdominal cavity (peritoneum) was light and unpigmented in C. carassius but was black with a pearly sheen in C. a. gibelio. Peritoneal colouring of the hybrids was variable – some individuals had darker and some had lighter colouring. The average number of gill rakers (Sp.br.) reached 48Á3 in diploid C. a. gibelio, 28Á6inC. carassius and 39Á4 in hybrids (Table I). This trait clearly distin- guished hybrids from parental species. Comparison of other meristic traits without Sp.br. discriminates C. a. gibelio from both C. carassius and hybrids, but C. carassius and hybrids could not be distinguished without Sp.br. (Table I). DCA of the number of gill rakers demonstrated clear differences between C. a. gibelio, C. carassius and their hybrids (Fig. 1). The number of gill rakers is useful for the identification of species and hy- brids of the genus Carassius (Berg, 1932). The average number of gill rakers found in diploid C. a. gibelio individuals is consistent with data published by other authors (Vasileva, 1990; Vetesˇnı´k, 2005). Only Kux (1982) found 41 gill rakers in a single hybrid of these species. This number is close to the average number (39Á4) Sp.br. of the hybrid individuals. The analysis of four microsatellite markers in control samples of C. a. gibelio and C. carassius produced alleles within or close to the size ranges described by Ha¨nfling et al. (2005); only MFW7 had an allele that was significantly different in size, but this allele was nevertheless consistent with alleles in C.
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