Relationships and genetic consequences of contrasting modes of speciation among endemic species of Robinsonia (Asteraceae, Senecioneae) of the Juan Fernández Archipelago, Chile, based on AFLPs and SSRs.

This study analyses and compares the genetic signatures of anagenetic and cladogenetic speciation in six species of the genus Robinsonia (Asteraceae, Senecioneae), endemic to the Juan Fernández Islands, Chile. Population genetic structure was analyzed by amplified fragment length polymorphism (AFLP) and microsatellite (simple sequence repeat, SSR) markers from 286 and 320 individuals, respectively, in 28 populations. Each species is genetically distinct. Previous hypotheses of classification among these species into subgenera and sections, via morphological, phytochemical, isozymic and internal transcribed spacer (ITS) data, have been confirmed, except that R. saxatilis appears to be related to R. gayana rather than R. evenia. Analysis of phylogenetic results and biogeographic context suggests that five of these species have originated by cladogenesis and adaptive radiation on the older Robinson Crusoe Island. The sixth species, R. masafuerae, restricted to the younger Alejandro Selkirk Island, is closely related to and an anagenetic derivative of R. evenia from Robinson Crusoe. Microsatellite and AFLP data reveal considerable genetic variation among the cladogenetically derived species of Robinsonia, but within each the genetic variation is lower, highlighting presumptive genetic isolation and rapid radiation. The anagenetically derived R. masafuerae harbors a level of genetic variation similar to that of its progenitor, R. evenia. This is the first direct comparison of the genetic consequences of anagenetic and cladogenetic speciation in plants of an oceanic archipelago.

Progressive migration and anagenesis in Drimys confertifolia of the Juan Fernández Archipelago, Chile.

A common mode of speciation in oceanic islands is by anagenesis, wherein an immigrant arrives and through time transforms by mutation, recombination, and drift into a morphologically and genetically distinct species, with the new species accumulating a high level of genetic diversity. We investigate speciation in Drimys confertifolia, endemic to the two major islands of the Juan Fernández Archipelago, Chile, to determine genetic consequences of anagenesis, to examine relationships among populations of D. confertifolia and the continental species D. winteri and D. andina, and to test probable migration routes between the major islands. Population genetic analyses were conducted using AFLPs and nuclear microsatellites of 421 individuals from 42 populations from the Juan Fernández islands and the continent. Drimys confertifolia shows a wide genetic variation within populations on both islands, and values of genetic diversity within populations are similar to those found within populations of the continental progenitor. The genetic results are compatible with the hypothesis of high levels of genetic variation accumulating within anagenetically derived species in oceanic islands, and with the concept of little or no geographical partitioning of this variation over the landscape. Analysis of the probability of migration within the archipelago confirms colonization from the older island, Robinson Crusoe, to the younger island Alejandro Selkirk.

Genetic variation (AFLPs and nuclear microsatellites) in two anagenetically derived endemic species of Myrceugenia (Myrtaceae) on the Juan Fernández Islands, Chile.

PREMISE OF THE STUDY: Anagenesis (or phyletic evolution) is one mode of speciation that occurs in the evolution of plants on oceanic islands. Of two endemic species on the Juan Fernández Islands (Chile), Myrceugenia fernandeziana and M. schulzei (Myrtaceae), believed to have originated anagenetically from different continental progenitors, the first is endemic to Robinson Crusoe Island and has no clear tie to continental relatives; the last is endemic to the younger island, Alejandro Selkirk Island, and has close affinity to M. colchaguensis in mainland Chile. METHODS: Using AFLPs and six nuclear microsatellites from 381 individuals representing 33 populations, we determined patterns of genetic variation within and among populations on both islands and between those of the islands and mainland. KEY RESULTS: Considerable genetic variation was found within populations on both islands. The level of gene diversity within M. schulzei was equivalent to that of its close continental relative M. colchaguensis. Genetic diversity was not partitioned geographically in M. fernandeziana and was weakly so and nonsignificantly in M. schulzei. CONCLUSIONS: The high genetic variation in both taxa is most likely due to anagenetic speciation. Subsidence of the older island Robinson Crusoe, landscape erosion, and restructuring of communities have severely reduced the overall island population to a single panmictic system. On the younger and less modified Alejandro Selkirk Island, slightly stronger patterns of genetic divergence are seen in M. schulzei. Because both species are genetically diverse and number in the thousands of individuals, neither is presently endangered in the archipelago.

Progenitor-derivative speciation in Pozoa (Apiaceae, Azorelloideae) of the southern Andes.

BACKGROUND AND AIMS: Studies examining patterns and processes of speciation in South America are fewer than in North America and Europe. One of the least well documented processes has been progenitor-derivative speciation. A particularly instructive example occurs in the southern Andes in the genus Pozoa (Apiaceae, Azorelloideae), which consists of only two diploid outcrossing species, the widespread P. coriacea and the geographically and ecologically restricted P. volcanica. This paper tests the hypothesis that the latter species originated from the former through local geographical and ecological isolation by progenitor-derivative speciation. METHODS: DNA sequences were analysed from Pozoa and the related South American genera Asteriscium, Eremocharis and Gymnophyton from non-coding regions of the plastid genome, ndhF-rpl32 and rpl32-trnL, plus incorporation of previously reported rpl16 intron and trnD-trnT intergenic spacer sequences. Amplified fragment length polymorphism (AFLP) data from 105 individuals in 21 populations throughout the entire range of distribution of the genus were used for estimation of genetic diversity, divergence and SplitsTree network analysis. Ecological factors, including habitat and associated species, were also examined. KEY RESULTS: Pozoa coriacea is more similar genetically to the outgroup genera, Asteriscium and Eremocharis, than is P. volcanica. At the population level, only P. volcanica is monophyletic, whereas P. coriacea is paraphyletic. Analyses of genetic differentiation among populations and genetic divergence and diversity of the species show highest values in P. coriacea and clear reductions in P. volcanica. Pozoa coriacea occurs in several types of high elevation habitats, whereas P. volcanica is found only in newly formed open volcanic ash zones. CONCLUSIONS: All facts support that Pozoa represents a good example of progenitor-derivative speciation in the Andes of southern South America.

Patterns of genetic diversity in colonizing plant species: Nassauvia lagascae var. lanata (Asteraceae: Mutisieae) on Volcan Lonquimay, Chile.

The effect of colonization on the distribution of genetic diversity within and among populations in relation to species characteristics remains an open empirical question. The objective of this study was to contrast genetic diversity within and among established and colonizing populations of Nassauvia lagascae var. lanata on Volcán Lonquimay (Araucanía Region, Chile), which erupted on 25 December 1988, and relate genetic diversity to biological characteristics of the populations. We analyzed a total of 240 individuals from 15 populations distributed along the Andes Cordillera using AFLP and obtained a total of 307 AFLP bands, of which 97.7% are polymorphic. Values of population differentiation (F(ST)) did not differ significantly among established and colonizing populations, but colonizing populations did have reduced levels of genetic divergence (as indicated by private and rare bands) and genetic variation (e.g., Shannon index). We conclude that a founder effect through limited numbers of founding propagules derived from nearby source populations has not yet been compensated for by subsequent population growth and migration. Low rates of secondary dispersal via running water, kin-structure within populations, and slow population growth seem to contribute to the slow recovery of genetic diversity.

Pleistocene refugia and polytopic replacement of diploids by tetraploids in the Patagonian and Subantarctic plant Hypochaeris incana (Asteraceae, Cichorieae).

We report the phylogeographic pattern of the Patagonian and Subantarctic plant Hypochaeris incana endemic to southeastern South America. We applied amplified fragment length polymorphism (AFLP) and chloroplast DNA (cpDNA) analysis to 28 and 32 populations, respectively, throughout its distributional range and assessed ploidy levels using flow cytometry. While cpDNA data suggest repeated or simultaneous parallel colonization of Patagonia and Tierra del Fuego by several haplotypes and/or hybridization, AFLPs reveal three clusters corresponding to geographic regions. The central and northern Patagonian clusters (approximately 38-51 degrees S), which are closer to the outgroup, contain mainly tetraploid, isolated and highly differentiated populations with low genetic diversity. To the contrary, the southern Patagonian and Fuegian cluster (approximately 51-55 degrees S) contains mainly diploid populations with high genetic diversity and connected by high levels of gene flow. The data suggest that H. incana originated at the diploid level in central or northern Patagonia, from where it migrated south. All three areas, northern, central and southern, have similar levels of rare and private AFLP bands, suggesting that all three served as refugia for H. incana during glacial times. In southern Patagonia and Tierra del Fuego, the species seems to have expanded its populational system in postglacial times, when the climate became warmer and more humid. In central and northern Patagonia, the populations seem to have become restricted to favourable sites with increasing temperature and decreasing moisture and there was a parallel replacement of diploids by tetraploids in local populations.

Characterization, genomic organization and chromosomal distribution of Ty1-copia retrotransposons in species of Hypochaeris (Asteraceae).

This study aims to analyze the diversity of Ty1-copia retrotransposons in 18 taxa of Hypochaeris, including two Old World species H. maculata (2n=2x=10) and H. angustifolia (2n=2x=8), and representatives of the South American species (16 accessions of 15 species; all 2n=2x=8). Analysis of 380 PCR-amplified sequences, corresponding to a conserved domain of the subset of Ty1-copia reverse transcriptase (rt) gene amplifiable with degenerate standard primers, showed high levels of intra- and interspecific heterogeneity. Nucleotide diversity (Pi) of the copia fragments was high in all species and varied from 0.229 (H. angustifolia) to 0.412 (H. chillensis). Higher sequence heterogeneity correlates positively with larger genome size among analyzed species. Phylogenetic analyses of amplified fragments revealed different patterns of intraspecific heterogeneity within species, with most sequences forming one well-supported main clade while a few sequences fall into small clades or are left ungrouped. The combined analysis of all sequences revealed the presence of three main clades and showed that highly diverged species contain closely related Tyl-copia group retrotransposons. One of the main clades includes rt sequences of all South American species and three sequences of their putative ancestor, H. angustifolia, but no sequence of the Old World H. maculata. FISH with copia retrotransposons in four Hypochaeris species, including H. maculata and H. angustifolia and New World H. apargioides and H. spathulata, revealed differences in the chromosomal distribution between the two groups. In Old World species copia retroelements are distributed over the whole length of the chromosomes, excluding rDNA sites and some centromeres. In the South American species the two largest chromosome pairs are enriched in copia, while most of the long arms of the two small pairs of chromosomes are devoid of these elements. The patterns of heterogeneity and chromosomal distribution of Ty1-copia retrotransposons in Hypochaeris are discussed in the context of the origin, genome evolution and organization of the South American species.

Karyotype diversification and evolution in diploid and polyploid South American Hypochaeris (Asteraceae) inferred from rDNA localization and genetic fingerprint data.

BACKGROUND AND AIMS: Changes in chromosome structure and number play an important role in plant evolution. A system well-suited to studying different modes of chromosome evolution is the genus Hypochaeris (Asteraceae) with its centre of species' diversity in South America. All South American species uniformly have a chromosome base number of x = 4 combined with variation in rDNA number and distribution, and a high frequency of polyploidy. The aim of this paper is to assess directions and mechanisms of karyotype evolution in South American species by interpreting both newly obtained and previous data concerning rDNA localization in a phylogenetic context. METHODS: Eleven Hypochaeris species from 18 populations were studied using fluorescence in situ hybridization (FISH) with 35S and 5S rDNA probes. A phylogenetic framework was established from neighbour-net analysis of amplified fragment length polymorphism (AFLP) fingerprint data. KEY RESULTS: A single 5S rDNA locus is invariably found on the short arm of chromosome 2. Using 35S rDNA loci, based on number (one or two) and localization (interstitial on the long arm of chromosome 2, but sometimes lacking, and terminal or interstitial on the short arm of chromosome 3, only very rarely lacking), seven karyotype groups can be distinguished; five of these include polyploids. Karyotype groups with more than one species do not form monophyletic groups. CONCLUSIONS: Early evolution of Hypochaeris in South America was characterized by considerable karyotype differentiation resulting from independent derivations from an ancestral karyotype. There was marked diversification with respect to the position and evolution of the 35S rDNA locus on chromosome 3, probably involving inversions and/or transpositions, and on chromosome 2 (rarely 3) concerning inactivation and loss. Among these different karyotype assemblages, the apargioides group and its derivatives constitute by far the majority of species.