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Spatial Structure of a Natural Mixed Topodeme of Subalpine Sorbus Taxa

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Spatial Structure of a Natural Mixed Topodeme of Subalpine Sorbus Taxa Vol. 77, No. 4: 305-311, 2008 ACTA SOCIETATIS BOTANICORUM POLONIAE 305 SPATIAL STRUCTURE OF A NATURAL MIXED TOPODEME OF SUBALPINE SORBUS TAXA DUAN GÖMÖRY, DIANA KRAJMEROVÁ Technical University in Zvolen, Faculty of Forestry TG Masaryka 24, SK-960 53 Zvolen, Slovakia e-mail: [email protected] (Received: August 6, 2007. Accepted: March 20, 2008) ABSTRACT Spatial distribution and genetic variation of a population of Sorbus chamaemespilus (L.) Crantz and putative hybrids between S. chamaemespilus, S. aria and S. aucuparia growing in the nature reserve Skalná Alpa (central Slovakia) were studied. The analysis of spatial patterns using Ripleys K-function revealed a significant clustering of the adults of both S. chamaemespilus and hybrid taxa at distances up to ~15 m and a strong affinity between both taxonomical groups, indicating similar ecological requirements. Bivariate point-pattern analysis considering cardinal direction showed that juvenile individuals of S. chamaemespilus are clustered around the adults up to the distance of ~2 m, whereas in hybrid taxa with larger and more dense crowns, juveniles are clustered at distances more than ~3 m from the adults. The analysis of genetic variation in a subset of adult shrubs using 4 nuclear mi- crosatellite loci revealed that unlike expected, there was no variation in S. chamaemespilus but several genotypes were found in the group of hybrid taxa. Implications for the reproduction system and conservation of the investi- gated taxa are discussed. KEY WORDS: Sorbus L., spatial structure, apomixis, nuclear microsatellites, dispersal. INTRODUCTION tal taxa. Generally, they are limited to the subalpine zone of high mountain ranges in the central part of Western Car- The genus Sorbus L., just like the whole family Rosace- pathians. Four taxa occur in relic dwarf pine stands of the ae, belongs to the most complicated plant taxa (Aldasoro et Ve¾ká Fatra Mts. in central Slovakia. Based on morpholo- al. 1998; Nelson-Jones et al. 2002). Flora Europaea lists gy, Bernátová and Májovský (2003) suggested the origin 113 Sorbus taxa of various ranks in Europe. Among them, from the hybridization of S. chamaemespilus and S. aria in there are five diploid species classified to different subge- the case of two taxa: S. zuzanae Bernátová et Májovský nera (Warburg and Kárpáti 1968): S. aucuparia L. (subgen. and S. haljamovae Bernátová et Májovský, whereas in the Sorbus), S. domestica L. (subgen. Cormus), S. torminalis cases of S. montisalpae Bernátová et Májovský and S. atri- (L.) Crantz (subgen. Torminaria), S. chamaemespilus (L.) montis Bernátová et Májovský, they suppose that these taxa Crantz (subgen. Chamaemespilus), and S. aria (L.) Crantz have originated from a complex hybridization process in- (subgen. Aria). These species are generally recognized and volving subgenera Sorbus, Aria and Chamaemespilus. Al- occupy relatively large ranges. Moreover, there is a plenty though the authors of the description did not explore the of local, mostly stenoendemic taxa of hybridogenous ori- reproduction system of this taxon in detail, they suggested gin. Most of them are supposed to have originated from the that it spreads clonally. hybridization between S. aria sensu lato and S. chamaeme- The mode of reproduction is reflected in spatial distribu- spilus, S. torminalis or one of the species of the subgenus tion. As documented by theoretical models, computer simu- Sorbus, with S. aria as the pollen donor. These hybrids are lations and empirical studies, the mechanisms of pollen generally triploid or tetraploid, and most of them are obli- flow, embryo formation, seed dispersal and vegetative spre- gatorily or facultatively apomictic (Jankun 1993; Campbell ad affect spatial patterns of the distribution of individuals, and Dickinson 1990; Nelson-Jones et al. 2002). distribution of offsprings in relation to the parents, and con- In Western Carpathians, several Sorbus microspecies ha- sequently also spatial genetic structures (Berg and Hamrick ve recently been described, among which six have pink or 1994; Burczyk et al. 1996; Hardy and Vekemans 1999; Shi- pink-bordered petals, indicating that S. chamaemespilus matani 2004). Naturally, dispersal-based spatial patterns are (the sole rosy-flowered Sorbus species) is one of the paren- subsequently modified by the influences of abiotic environ- 306 SPATIAL STRUCTURE AND GENETIC VARIATION OF A SORBUS CHAMAEMESPILUS POPULATION Gömöry D. et al ment and biotic interactions, including intraspecific interac- followed by 35 amplification cycles (denaturation 45 s at tions among different age classes (Caspersen and Saprunoff 95°C, annealing 45 s at 55°C, extension 60 s at 72°C) and 2005; LePage et al. 2000; Dovèiak et al. 2008). a final extension step for 5 min at 72°C. Before loading The objective of this study was to identify spatial distri- amplification products were diluted with the equal amount bution patterns in subalpine Sorbus taxa and to make infe- of loading dye consisting of 0.5% bromophenol blue, 8 rence on the reproduction system of the respective taxon M urea and 10 mM sodium hydroxide and denatured at based on spatial patterns and genetic variation of the re- 95°C for 3 minutes and placed on ice immediately. Sam- spective taxon. ples were loaded on 6% polyacrylamide sequencing gels, run at constant power 40 W for about 5 hours and bands were visualised by silver staning (Bassam et al. 1991). MATERIALS AND METHODS After fixing, gels were kept at 4°C overnight, and then air dried and scanned. All investigations were performed in the National Nature Spatial analyses were performed separately for S. chama- Reserve Skalná Alpa in central Slovakia. The reserve, cove- emespilus and jointly for all hybrid taxa. Spatial distribu- ring 524 hectares, extends over several vegetation zones tion patterns were analyzed using the Ripleys K-function: from 940 to 1463 m a.s.l. Plant communities in the summit -2 -1 part, growing on Triassic dolomites, belong to ass. Adeno- K(t) = n A Si Si¹j wij It(uij), stylo-Pinetum mughii and are formed mainly by shrubs of mountain dwarf pine and rowan with scattered Norway spru- where t is the distance for which K is calculated, n is the ce. In addition to S. aucuparia, six Sorbus taxa have been number of Sorbus individuals in the analyzed plot, A is the found here dispersed in dwarf pine stands, namely S. chama- area of the plot, uij is the distance between the individuals emespilus, S. aria, S. zuzanae, S. haljamovae, S. montisalpae i and j, It(uij) is a counter variable (It is 1 if uij < t and 0 and S. atrimontis (Bernátová and Májovský 2003). otherwise), and wij is a weighting factor for edge correction All shrubs of Sorbus chamaemespilus and putative hy- (Ripley 1976; Haase 1995). In the case of a random Pois- brids were mapped within a 70×50 m rectangular area aro- son distribution of individuals, the expected value of K(t) = und the summit of Skalná Alpa (longer side oriented along pt2. The results are reported as the variance-stabilizing squ- the crest). One S. aria, 30 adult S. chamaemespilus, 35 ju- are-root transformation of K(t) as: venile S. chamaemespilus, 10 S. haljamovae, 19 S. zuza- L(t) =K/p t, nae, 4 S. montisalpae and 35 juvenile hybrid individuals Ö t (which could not be classified to taxa without ambiguity) were found on the mapped area. Positions of individuals where L(t) equals zero when the dispersion pattern is (estimated as the center of the respective rooting area of random at the scale of the radial distance t, and L(t) > 0 in- the shrub) were measured using the Trimble GeoXT geo- dicates aggregation, whereas L(t) < 0 indicates a uniform graphic positioning system with the precision of ±0.3 m, pattern. supplemented by tape measurements. As hybrids form lar- In the bivariate case (comparison adults vs. juveniles), ge shrubs with asymmetric crowns, the position of an indi- the formula for the K-function becomes: vidual need not be identical with the centre of the crown -1 -1 projection. Polycormons were recognized as discrete indi- K12(t) = n1n2 A Si Sj wij It(uij); i ¹ j, viduals if there was no visible connection (e.g., through la- yered branch) with another individual. n1 being the number of adults, n2 the number of juveni- Leaves were collected from a subset of flowering (unam- les, and uij the length of the vector from the ith adult indi- biguously identified) Sorbus shrubs in June 2006. In total, vidual to the jth juvenile. 1 S. aria, 29 S. chamaemespilus, and 18 hybrids (4 S. hal- To account for potential anisotropies in the distribution jamovae, 11 S. zuzanae and 3 S. montisalpae) were sam- of juveniles around the adults because of slope effect, we pled. Leaves were placed in Ziploc bags with silica gel and also performed bivariate analysis considering the cardinal stored at room temperature. To extract genomic DNA, two direction for individuals growing at the NW-facing (steep) leaf discs in diameter of 8 mm were cut and ground to a fine slope: powder in a mixer mill using tungsted beads. Genomic -1 -1 DNA was extracted using the CTAB method according to Kq12(t) = n1n2 A Si Sj wij It(q)(uij); i ¹ j, Doyle and Doyle (1987), which was modified for a smaller amount of starting material. For genotyping, microsatellite where Kq12(t) is the K-function value for the respective markers MSS1, MSS3, MSS4 and MSS10 according to cardinal direction (defined as the direction of quadrant axes Oddou-Muratorio et al. (2001) were used. The choice of of circles centered at the adults, whereby in our case, the markers was based on a preliminary study with S. aria × S. axes were oriented parallel and/or perpendicular to contour chamaemespilus hybrids (unpublished data).
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