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<I>Corbicula Fluminalis</I>

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<I>Corbicula Fluminalis</I> Folia biologica (Kraków), vol. 57 (2009), No 3-4 doi:10.3409/fb57_3-4.177-185 CytogeneticandMorphologicalCharacterizationof Corbiculafluminalis (O.F.Müller,1774) (Bivalvia:Veneroida:Corbiculidae): TaxonomicStatusAssessmentofaFreshwaterClam LidiaSKUZA,AnnaMaria£ABÊCKA andJózefDOMAGA£A Accepted April 20, 2009 SKUZA L., £ABÊCKA A. M., DOMAGA£A J. 2009. Cytogenetic and morphological characterization of Corbicula fluminalis (O.F.Müller,1774)(Bivalvia:Veneroida:Corbiculidae): taxonomic status assessment of a freshwater clam. Folia biol. (Kraków) 57: 177-185. Chromosomes of Corbicula fluminalis were characterized by karyotype analysis and nucleolar organizer region (NORs) localization. The triploid chromosome number was confirmed as 54; the karyotype is composed of 3 metacentric, 15 submetacentric and 36 subtelo-acrocentic chromosomes. Silver staining revealed nucleolar organizers on the telomeric regions of three subtelo-acrocentic chromosomes. This is the first study on chromosomes of C. fluminalis. The results are discussed with regards to Corbicula species as well as its relationships to other mollusc species based on cytogenetic characters and morphometric of the shells. Key words: AgNOR, chromosomes, cooling water, Corbicula fluminalis, invasive species, karyotype, triploid. Lidia SKUZA, University of Szczecin, Department of Cell Biology, W¹ska 13, Szczecin 71-415, Poland. E-mail: [email protected] Anna Maria £ABÊCKA, Józef DOMAGA£A, University of Szczecin, Department of General Zo- ology, Felczaka 3c, Szczecin 71-412, Poland. E-mail: [email protected], [email protected] [email protected] Taxonomy of Corbiculidae is not well resolved. invasion of this clam in the Danube River, in Ger- A revision of the 50 species described by HEUDE many, used the name Corbicula fluminea/fluminalis. (1880) has shown that as many as 43 of them are HAESLOOP (1992), but also MORTON (1982) finally synonyms of Corbicula fluminea (O. F. Müller, described C. fluminalis as Corbicula cf. fluminalis, 1774) (PRASHAD 1929). From among many Cor- pointing to its uncertain systematic classification. bicula species earlier described from Asia, MOR- A molecular investigation of the Corbicula TON (1982) distinguished only two: C. fluminea populations sympatrically occurring in the Rhine (freshwater) and Corbicula fluminalis (O. F. Mül- River has revealed the co-existence of two evolu- ler, 1774) (estuarine). In the latest report on the tionarily different lineages producing cryptic hy- malacofauna of Germany, C. fluminea and C. flu- brids (PFENNINGER et al. 2002). Taking into minalis are also mentioned as two species (GLÖER regard their potential incomplete reproductive iso- &ZETTLER 2005). They are distinguished on the lation, PFENNINGER et al. (2002) have proposed basis of the structure of the shell, environmental that different lineages of Corbicula can be treated preferences and mode of reproduction (MORTON rather as the initial stage of a group of species than 1982; BIJ DE VAATE &GREIJDANUS-KLAAS 1990; as a few well-defined species. RAJAGOPAL et al. 2000). However, the distinction of species in the genus of Corbicula remains un- In view of the above issues, it is desirable to certain and questionable. The problem is the atypical search for new features that could be used in the sympatric appearance of the invasive C. fluminalis taxonomy of Corbicula. Chromosomal knowl- accompanied by C. fluminea in Europe (CSÁNYI edge is increasingly recognized as an important 1998-1999; REUMER 1993; SWINNEN et al. 1998; force in animal evolution and therefore the aim of NGUYEN & DE PAUW 2002; DOMAGA£A et al. this study was to describe the karyotype and iden- 2004; £ABÊCKA et al. 2005; PAUNOVIÆ et al. 2007). tify the active organizer regions, NORs, in C. flu- TITTIZER &TAXACHER (1997) in a description of the minalis chromosomes. 178 L.SKUZA etal. Material and Methods drop was withdrawn back into the Pasteur pipette (BOROÑ et al. 2004). The C. fluminalis specimens studied were col- The slides were stained with a 2% orcein solu- lected from a channel discharging cooling water from tion at 46oC for 30 minutes. Observations were the Power Plant Dolna Odra (N 53°12’ E 14°27’, conducted under a Nicon Eclipse E600 microscope Western Pomerania Region, N-W Poland). The with an immersion objective of 100 H magnifica- clams were collected manually by free diving, at a tion. The photographs were stored in computer depth of 0.5-2.0 m between May and July 2007. memory using an Applied Imaging ER-3339 cam- The grain size of the bottom substrate and particle era (Applied Imaging International Ltd.). In order sorting was measured by a laser Malvern Master- to detect the active NORs, the AgNOR chromo- sizer Micro Ver. 2.19 in order to determine the somes were stained according to the method de- type of river bottom in which the clams lived. Wa- scribed by HOWELL &BLACK (1980). ter parameters such as temperature (°C), dissolved -3 oxygen (O2 dm ), pH and electrolytic conductiv- Morphometric measurements of the chromosomes ity (FS cm-1)weremeasuredbyanElmetronmeas- were made using the Micromeasure Program Ver. 3.3 uring device. (www.colostate.edu/Depts/Biology/MicroMeasure). The lengths of the chromosome arms were meas- After transportation to the laboratory, the clams ured (short arm, p; long arm, q). The relative chro- were placed in aerated aquariums. The shell di- mosomelength(RL),theratioofthelengthsofthe mensions (length L, height H and width W) were arms (q/p) and the centromeric index (CI) were measured by an electronic Etalon slide-calliper to calculated: an accuracy of 0.1 mm (Figs 1A, B). The elonga- tion index (L/H) H 100, the height index (H/L) H 100 RL = 100 H [(p+q)/ total haploid length)] and the width index (W/H) H 100 were calculated. Detailed shell morphology was observed under a H Carl Zeiss Stemi 2000-C microscope, under a CI = 100 [p/(p+q)] magnification of 50 H. The systematic position of The chromosomes were classified according to the C. fluminalis specimens was established on the the criteria proposed by LEVAN et al. (1964). The basis of the key for identification of freshwater karyogram was made with the help of the Genus molluscs (GLÖER &MEIER-BROOK 1998). Voucher Program Ver. 3.1 (Applied Imaging International specimens of the clams studied and their shells are Ltd.). stored at the Department of General Zoology, Uni- versity of Szczecin (acronym KZOUSZ, Katedra Zoologii Ogólnej, Uniwersytet Szczeciñski). Results Cytogenetic investigation was performed on the basis of analysis of 26 metaphase plates isolated Theclamslivedonasandyandmuddyriverbot- from 20 specimens of C. fluminalis. Because of a tom of moderately well sorted grains. The size of low level of the mitotic index, in vivo cell division grains and their percent contributions encom- was stimulated by a 0.4% solution of cobalt chlo- passed: medium sand 19.4%, fine sand 64.5%, ride was applied in the amount of 0.05 ml, injected veryfindsand11.9%,verycoarsesilt0.9%,coarse directly into the body of each specimen. Cobalt silt 3.0%, medium silt 0.3%. chloride blocks two major steps of cellular respira- tion, inducing tissue hypoxia, which leads to cell Themeanwatertemperatureduringtheperiodof study was 25.3°C (SD = 4.3), pH 8.8 (SD = 0.18), proliferation (WONICKI 2004).After60hoursthe -3 dissolved oxygen 11.76 mg O2 dm (SD = 0.9), clams were injected in vivo with 0.1% solution of -1 colchicine in the amount of 0.25 ml per individual and electrolytic conductivity 863.7 FS cm (SD = per 6 hours. After this time the gills were dissected 95.7). and hypotonized in a 0.01% NaCl solution for 1 The shells of C. fluminalis were oval-triangular, hour. The suspension was centrifuged (1000 rpm, strongly convex and of clear asymmetry (Figs 1A, B). 10 min) in a MPW-350R centrifuge (MPW Med. The shell umbo was above the ligament, slightly Instrument),theprecipitatewasfixedina3:1solu- rotated and directed to the front of the shell. The tion of methanol and glacial acetic acid and centri- periostracum was glossy, olive-green coloured fuged again at 1000 rpm for 10 minutes. The and covered with ribs. The number of ribs per 10 supernatant was replaced by fresh fixative solu- mm of the shell surface was 12-16. The en- tion. The centrifugation was repeated twice and dostracum was intensely violet in the ventral, pos- then the cellular suspension was suspended in a terior and anterior parts, while in the middle part 50% acetic acid solution. This cell suspension was and under the umbo it was white-violet with or- dropped onto a microscope slide, heated to angespots.Thepalliallinewasclearlymarked,the 40-45oC and after a few seconds the suspension adductor muscle impressions were well visible. Corbiculafluminalis ChromosomesandShell 179 A H L W B Fig. 1. The shells of C. fluminalis. (A) Shell shape, endostracum and periostracum. (B) Umbo view. Abbreviations: L – length, H–height,W–width.Bar=1cm. The morphometric data concerning the shells of into 18 groups of 3 phenotypically similar chromo- the individuals used in the cytogenetic studies are somes(Fig.2B).Analysisofthecentromericindex presented in Table 1. Analysis of the number of (Table 2) has proved that the karyotype is com- chromosomes in 26 metaphase plates revealed that the karyotype of the studied species is composed posed of 3 metacentric, 15 submetacentric and 36 of 54 chromosomes, (Fig. 2A) that can be divided subtelo-acrocentric chromosomes (Fig. 2B). 180 L.SKUZA etal. Table 1 Morphometric data of shells of C. fluminalis from the Odra River, Poland L(mm) W(mm) H(mm) (L/H)H100 (H/L)H100 (W/H)H100 mean 19.0 15.9 19.1 100.1 99.9 82.8 SD 3.6 3.4 3.9 3.6 3.5 2.3 min 10.3 7.6 9.8 95.5 91.2 77.6 max 22.6 19.3 22.8 109.5 104.6 87.9 Abbreviations:L–shelllength,H–shellheight,W–shellwidth.
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