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Allozyme Variation and Possible Phylogenetic Implications in Abies Cephalonica Loudon and Some Related Eastern Mediterranean Firs

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Allozyme Variation and Possible Phylogenetic Implications in Abies Cephalonica Loudon and Some Related Eastern Mediterranean Firs Allozyme Variation and Possible Phylogenetic Implications in Abies cephalonica Loudon and Some Related Eastern Mediterranean Firs By B. Fady1) and M. T. Conkle2) (Received 4th August 1993) Summary northwards, the species is progressively replaced by A. A total of 22 loci were assayed in several populations of borisii regis which becomes the only fir species in northern Abies cephalonica, A. borisii regis, A. bornmuelleriana and Greece. These species are valuable for reforestation in A. alba using horizontal starch gel electrophoresis. Wdthin- Mediterranean France both from an ecological and a and between-population diversity were analyzed as well as silvicultural perspective. A. cephalonica has a large and between-species diversity. Mean expected heterozygosity scattered geographic distribution (Panetsos, 1975) under a was high and ranged between 0.175 and 0.290. The per wide range of ecological conditions contributing to isola centage of loci polymorphic was also very high, from 52.9 tion, hybridization and introgression phenomena. to 87.5. All populations except the A- borisii regis prove nance were either at equilibrium or significantly deficient in Terpene analyses (Koedam, 1981; Mitsopoulos and Panet heterozygotes, which suggests a certain level of inbreeding. sos, 1987; Fady et al., 1992) tend to prove that A. cephalo Some alleles were species-specific at the following loci: nica has a highly variable genetic structure and emerged Pgil, Pgi2, Lap, Got2 and Mnrl. Most of the genetic diver from an ancestral species common to A. alba and A. gence between the studied species and provenances appea bornmuelleriana during the Pleistocene (Fady et al., 1992). red recently, during and after the last glaciation. The only known electrophoretic study (Scaltsoyiannes et Key words: Abies cephalonica, allozymes, genetic diversity, phylo- al., 1991) also indicated that genetic variability in A. geny, Abies alba, Abies bornmuelleriana, Abies borisii cephalonica is high, based on a limited number of enzyme regis. systems. The aim of this study was to quantify genetic diversity Introduction in A. cephalonica aad to provide additional information on Abies cephalonica is a fir species endemic to the moun genetic variation* patterns and probable phylogenetic tains of central and southern Greece. From central Greece history of Abies in the eastern Mediterranean. *) INRA, Unite Experimental d'Amelioration des Arbres For- estiers Mediterraneens, Domaine du Ruscas, 4935 Route du Materials and Methods Dom, 83237 Bormes-Les-Mimosas cedex, France Wind-pollinated seeds were collected from 5 provenan *) Institute of Forest Genetics, Pacific Southwest Research Station, USDA Forest Service, P.O. Box 245, Berkeley, CA 94701, USA ces of A. cephalonica and 1 of A. borisii regis in Greece, Table 1. — Description of sampled material. Provenances coordinates species mountain range Pril 37°16N,22°18E A. cephalonica Taygetos (Greece) Kolo 38°33 N, 22°29 E A. cephalonica Parnassos (Greece) Veti 37°33 N, 22°15 E A. cephalonica Menalon (Greece) Ceph 38°14 N, 20°32 E A. cephalonica Cephallenia island (Greece) Evia 38°40 N, 23°30 E A. cephalonica Eubboea island (Greece) Hybr 39°30 N, 21°30 E A. borisii regis Pindos (Greece) Kzlk 41°10N, 33°50E A. bornmuelleriana Kastamonu (Turkey) Bald 41°05 N, 33°50 E A. bornmuelleriana Kastamonu (Turkey) Alba 48°28 N. 6°58 E A. alba Vosges (France) Sllvae Genetica 42, 6 (1993) 351 2 provenances of A. bornmuelleriana in Turkey and 1 composition (Fady et al., 1991). Both embryo and megaga- provenance of A. alba in France (Table 1). The species' metophyte tissues were used for the isozyme analysis. natural distributions are shown in figure 1. Samples were Extraction and horizontal starch gel electrophoretic proce made of bulked seeds using 30 trees per provenance for dures were as described in Conkle et al. (1982) and Fady all populations except A. borisii regis. In A. borisii regis, and Conkle (1992). parent identities were maintained: 403 seeds were assayed A total of 22 loci from 16 enzyme systems were scored: to calculate allele frequencies for 19 mother-trees. The acid phosphatase (ACP, EC 3.1.3.2), aconitase (ACO, EC A. cephalonica sample included provenances representative 4.2.1.3), catalase (CAT, EC 1.11.1.6), glutamate dehydro- of the geographic variability found for other traits such genase (GDH, EC 1.4.1.3), glutamate oxaloacetate trans- as morphology, height growth, phenology and terpene aminase (GOT, EC 2.6.1.1), glutathione reductase (GR, EC 352 Table 2. — Genetic diversity for 17 allozyme loci (14 loci for A. borisii regis) in 9 Abies provenances. Species means are given lor 22 loci. Standard error in parentheses. Provenance Mean Mean number Percentage Mean heterozygosity Mean FjS sample size of alleles of loci observed - expected(t>) per locus per locus polymorphic(a) Ceph 19.6(3.3) 1.9(0.2) 70.6 0.207 (0.056) 0.290 (0.062) 0.286 Evia 24.2 (3.2) 1.9(0.2) 70.6 0.225 (0.063) 0.250 (0.064) 0.100 Kolo 28.5 (3.6) 1.7(0.1) 64.7 0.140 (0.048) 0.242 (0.059) 0.421 Pril 23.2 (2.2) 1.9(0.2) 70.6 0.200 (0.048) 0.223 (0.056) 0.103 Veti 18.1 (3.1) 1.7(0.2) 52.9 0.147 (0.062) 0.192(0.060) 0.234 Alba 37.4 (3.9) 1.6(0.1) 58.8 0.174 (0.054) 0.209 (0.059) 0.167 Bald 29.6 (2.8) 1.7(0.1) 64.7 0.179 (0.057) 0.227 (0.055) 0.211 Kzlk 11.9(0.6) 1.6(0.1) 52.9 0.163 (0.045) 0.175(0.049) 0.069 Hybr 18.8(0.1) 2.1 (0.2) 87.5 0.313 (0.068) 0.279 (0.057) -0.122 A. cephal.(c) 96.0 (13.5) 2.0 (0.2) 72.7 0.161 (0.040) 0.221 (0.052) 0.271 (Fit) A. bornmu (c) 39.5 (2.9) 1.8(0.2) 59.1 0.161 (0.044) 0.198(0.047) 0.187 (Fjt) A. alba 35.6 (3.4) 1.6(0.1) 54.5 0.149 (0.044) 0. 182(0.048) 0,181 la) a locus is considered polymorphic if more than one allele was detected. (b) unbiased estimate (Nei, 1978). (c) estimate using pooled data without Considering population subdivisions. Table 3. — Fixation index (F.g) and probably of deviation from Hardy-Weinberg equilibrium estimated using x* test between expected and observed allele frequencies at polymorphic loci. Significant deviations (p < 0.05) indicate that F.g is significantly different from zero. 3a- Species estimates (pooled data without considering population subdivisions) Locus A. cephalonica A. alba A. bornmuelleriana A borisii regis Fis P Fis P F* P Fis P Aco .605 .000 .392 .024 .384 .049 -.084 .743 Acp2 -.014 1.000 -.010 .893 .822 .003 -.056 .866 Got1 -.053 .639 - - - -.027 1.000 Got2 .365 .028 .085 .563 -.012 1.000 -.161 .545 Got3 -.055 .812 .447 .016 -.072 .887 -.162 .833 Gr .584 .000 .703 .000 .227 .166 .200 .331 Idh -.099 .407 -.016 .970 -.137 .313 nd nd Lap .441 .000 .109 .382 .619 .000 -.267 .282 Mdh2 -.013 .873 -.011 1.000 nd nd Mnr1 .259 .003 -.159 .339 .235 .077 -.080 .819 Mnr2 - . -.056 .866 Pep .447 .023 -.333 .315 -.157 .104 nd nd 6-Pgd -.022 .794 .256 .088 -.119 .456 -.056 .866 Pgi1 -.065 .586 - - -.119 .441 -.512 .088 Pg\2 .067 .285 - - -.016 1.000 -.250 .543 Pgm .660 .000 - - - nd nd Skdh -.051 .984 - -.025 .910 nd nd Ugpp - - 1.000 .000 - - nd nd 3b- A. cephalonica provenance estimates Locus Ceph Evia Kolo Pril Veti Fis P Fis P Pis p F* P Ffe P Aco .333 .346 .455 .346 1.00 .002 .200 .227 Acp2 - - -.053 1.00 - - Got1 -.167 .595 -.061 .801 -.029 1.00 -.048 1.00 Gr .784 .000 .649 .003 .357 .078 .267 .174 .792 .000 Idh -.600 .343 -.190 .456 .308 .127 -.159 .487 -.538 .120 Lap .451 .017 .220 .189 .793 .000 -.091 .703 -.050 .873 Mdh2 -.014 1.00 -.011 1.00 -.021 .918 - -.014 1.00 Mnr1 .596 .004 -.117 .609 .165 .309 .184 .263 .286 .159 Pep .491 .211 .083 .655 . 1.00 .011 .100 .633 1.00 .021 6-Pgd . -.014 1.00 -.043 .832 -.048 .825 - - Pgi1 .065 .635 -.200 .236 -.297 .058 -.067 .782 .093 .697 Pgi2 .042 .529 -.257 .162 .237 .027 -.078 .977 -.263 .562 Pgm 1.000 .000 -.032 1.00 - - - - - - Skdh -.143 1.00 -.125 .796 - - -.069 .992 - - —: locus was monomorphic for the population, rtd: no data 353 1.6.4.2), isocitrate dehydragenase (IDH, EC 1.1.1.42), leucine mate dehydrogenase (SkDH, EC 1.1.1.25), UDP-glucose amino peptidase (LAP, EC 3.4.11.1), malate dehydrogenase pyrophosphorylase (UGPP, EC 2.7.7.9). (MDH, EC 1.1.1.37) menadione reductase (MNR, EC 1.6.99.2), Allele frequencies for all 22 loci are given only for .peptidase (PEP, EC 3.4.11.1), phosphoglucomutase (PGM, species (pooled frequencies without considering population EC 2.7.5.1), 6-phosphogluconic dehydrogenase (6-PGD, EC subdivisions). Comparisons between A. cephalonica prove 1.1.1.44), phosphoglucose isomerase (PGI, EC 5.3.1.9), shiki- nances were made using 17 loci (see Table 4) and allele Table 4. — Estimated allele frequencies for 17 allozyme loci and heterogeneity of allele fre quencies for polmorphicf loci (0.95 criterion) between Abies cephalonica provenances, (n) is the sample size per locus. Abies cephalonica provenances heterogeneity Locus Ceph Evia Kolo Pril Veti df prob.
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