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Patterns of Genetic Variability in Populations of Adenostyles Cass Delpinoa, n.s. 44: 103-114. 2002 Patterns of genetic variability in populations of Adenostyles Cass. complex (Asteraceae) along the Apennine chain 1 1 1 ANNA MARIA PALERMO ,GIUSEPPE PELLEGRINO ,MARIA ELENA NOCE , 2 1 LILIANA BERNARDO ,ALDO MUSACCHIO 1Dipartimento di Ecologia, 2Orto Botanico, Università della Calabria, I-87036 Arcavacata di Rende, CS, Italy. Riassunto. In popolazioni disgiunte di Abstract. Allozymes and RAPDs markers were Adenostyles presenti lungo l'Appennino, catena tested in disjunct populations of Adenostyles com- montuosa che rappresenta un buon modello di plex along the Apennine chain, which is a good gradiente eco-geografico, sono stati testati allo- eco-geographic north-south gradient model. In our zimi e RAPD. I RAPD sono risultati più variabili survey, RAPD was higher than allozyme variability, rispetto agli allozimi, giacché ogni individuo ha with each individual showing a different RAPD mostrato un genotipo differente dagli altri. genotype. Both kinds of markers pointed out that Entrambi i marcatori hanno dimostrato che la majority of total genetic diversity resides within diversità genetica maggiore è presente all'interno populations, while geographic groups are scarcely delle popolazioni, mentre i gruppi geografici divergent and no clear geographic pattern was sono scarsamente divergenti e non è stato riscon- detected. Indeed, the dendrograms indicate that trato alcun gradiente geografico. La struttura populations group together without congruence genica spaziale non sarebbe da attribuire al flus- with their regional location. Data tend to exclude so genico, ma piuttosto rappresenterebbe la con- that this spatial genetic structure may be attributed seguenza di contatti secondari. Questi eventi pro- to a constantly maintained gene flow, but rather babilmente avvennero durante i cambiamenti cli- could be explained as a consequence of secondary matici del Quaternario, mentre la separazione fra contact events. These events probably occurred le popolazioni esaminate sarebbe avvenuta di during the climatic changes of Quaternary, and the recente. In aggiunta, sono stati discussi gli aspet- separation of the examined populations seems very ti conservazionistici, considerando sia la bassa recent. A conservational point of view is discussed, consistenza delle popolazioni sia la riduzione dei considering the low number of individuals in these loro ambienti. populations and the scarcity of their habitats. Key words: Adenostyles, allozymes, Apennine chain, Genetic variability, RAPDs, Spatial genetic structure INTRODUCTION vivor populations on the Alps, few studies have been conducted on the behaviour of In these last years, with the develop- herbaceous species during glaciations in ment of molecular markers, greater atten- the Mediterranean basin (COMES & tion has been placed to phylogeographic KADEREIT, 1998), which has already pro- studies on plants. In particularly, refugia ven to be an important refugium area for and recolonization of arctic and alpine taxa many tree species (COMES &KADEREIT, have been investigated (i.e., BAUERT et al., 1998; WILLIS, 1996). In this context the 1998; STEHLIK et al., 2001), with the aim Apennine chain, which has been viewed as of substantiating or disproving the nuna- an useful model of eco-geographic north- takker theory (STEHLIK, 2000). south gradient, represents a potential refu- Independently from the occurrence of sur- gium for alpine species and, eventually, a secondary colonisation route. In fact, its ties of an unique species, namely A. alpina orophilous floristic component includes, (FIORI, 1925-29), whereas in more recent among others, several Alpine species, time they have been treated discordantly as capable of reaching different latitudes and different subspecies or species (PIGNATTI, that have spread along the chain presu- 1982; WAGENITZ, 1983). In particular for mably during the Quaternary climatic these territories A. glabra (Mill.) DC., A. change. Due both to the peculiar orogra- australis (Ten.) Nyman and A. orientalis phic conformation and to the anthropic Boiss. have been reported, although their pressure, the populations of these species, differentiation has been based on few living in disjunction with respect to their highly variable and quantitative characters. main areas of distribution, are often Infact, these species are notoriously diffi- strongly isolated and progressively rare- cult to determine and some reports of their fied (PIGNATTI, 1980; PASSALACQUA & presence are questioned (PIGNATTI, 1982). BERNARDO, 1997). It has been shown that populations from The study of spatial structure of these central and northern zones of Apennines species with genetic markers may contri- were erroneously referred to A. glabra, bute to elucidate their evolutionary history whereas they belong to A. australis. On the and to shed light on the ecological and contrary, the alpine populations are refer- genetical influences of the colonisation red to A. glabra and not to A. australis. and isolation events experienced by these (PIGNATTI, 1982). In any case, regardless of taxa (THOMPSON, 1999; FREVILLE et al., taxonomic implications Apenninic 2001). Moreover, recent occurrence of Adenostyles complex may represent a good bottleneck or expansion in range may have model for exploring the pattern of genetic significant effects on actual population variation within and among populations of genetic structure (HANNAN &ORICK, an Alpine element, progressively further 2000). Unfortunately, there is a lack of from its main distribution area. As a matter interest towards the genetic structure of of fact, its effectively linear scattering natural plant populations along the transet along a north-south environmental gra- of this region (ZHANG et al., 2001; dient may reveal patterns of gene flow or GRÜNANGER et al., 1998; FRIZZI & phylogeografic routes beside their taxono- TAMMARO, 1991). mic evaluation. Consequently, in this paper The genus Adenostyles Cass. is preva- we chose to treat all examined populations lently distributed throughout the Alpine as belonging to Adenostyles complex. territories, but its range extends also to In addition, because no population Northern Spain, Corsica (France), Greece genetic information exists on these spe- and along the entire Apennine chain in cies, genetic variability analysis may Italy (including mountains of Calabria and encourage future conservation measures. Sicily) (WAGENITZ, 1983; TUTIN, 1976). Knowledge of population genetic structure The Apenninic entities of Adenostyles, provides an historical perspective of evolu- which occupy wet and shady habitats in tionary changes that characterise a species beech forests above 1400 m a.s.l., repre- and allows to predict how populations will sent a complex of strictly related taxa, respond to future natural events similar to each other from both a morpho- (VRIJENHOEK, 1987; HUENNEKE, 1991; logical and ecological points of view. In LUTZ et al., 2000). Consequently, the suc- the past, they have been regarded as varie- cessful preservation of populations of rare, 104 endangered or fragmented taxa may greatly benefit from similar genetic varia- tion results. In this paper we investigate the allozy- mic and RAPD variability of populations of the Apenninic Adenostyles entities. These markers have been widely applied to describe the genetic structure and diver- gence existing within and among popula- tions (HAMRICK &GODT, 1989; EDWARDS, 1998), with the aim of understanding the evolutionary factors responsible for their variation (SUN et al., 1999). RAPDs allow to emphasise variability in populations of endemic or rare plant species, even when allozymes failed in this goal (AYRES & RYAN, 1999). In brief, the major objectives of the pre- sent study are a) to evaluate relative levels Fig. 1 - Geographic distribution of populations of of genetic variation within Apenninic Adenostyles complex along the Apennine populations, b) to quantify differentiation chain among populations and gene flow, c) to scattered populations and the relative scar- assess genetic distances between popula- city of their habitats (CORBETTA et al., tions in relation to their geographic distan- 1998). ces, d) to indicate the potential consequen- ces of disjunction and the causes responsi- ble for the present Adenostyles distribu- MATERIALS AND METHODS tion, e) to determine the possible conserva- tional implications, since Adenostyles may Plant material be included at least among the rare species, Samples were collected from two popu- considering the low consistency of their lations in alpine localities and from eight Tab. 1 - Sampled populations of Adenostyles Sites Region Abbr. N. of sampled analyzed Allozyme sRAPDs Cozzo del Pellegrino Pollino massif (northern Calabria) PO-1 15 8 Tavolara Pollino massif (northern Calabria) PO-2 15 8 River Pollino massif (northern Calabria) PO-3 15 8 Botte Donato Sila (central Calabria) SI-1 11 8 Fontana Colonnello Sila (central Calabria) SI-2 66 Cervati Cilento (Campania) CI-1 15 8 Sicignano Cilento (Campania) CI-2 15 8 Mount Velino Marsica mountains (Abruzzo) AB-1 11 8 Magasa Eastern Alps (Lombardia) AL-1 15 8 Trento Eastern Alps (Trentino) AL-2 44 105 populations along the Apennines chain mic number of all Adenostyles is constant (Fig. 1). All populations were constituted throughout the genus (2n=38) (FEDEROV, by a few specimens, but in all cases the 1974). numbers of studied samples were represen- tative (Tab. 1). One or two leaves were col- RAPDs lected from each of individuals per popula-
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