Gene transfer across species boundaries in bryophytes: evidence from major life cycle stages in Homalothecium lutescens and H. sericeum.

BACKGROUND AND AIMS: The mosses Homalothecium lutescens and H. sericeum are genetically, morphologically and ecologically differentiated; mixed populations sometimes occur. In sympatric populations, intermediate character states among gametophytes and sporophytes have been observed, suggesting hybridization and introgression in such populations. METHODS: We determined genotypes using bi-allelic co-dominant single nucleotide polymorphism (SNP) markers, specific to either H. lutescens or H. sericeum, to estimate the degree of genetic mixing in 449 moss samples collected from seven sympatric and five allopatric populations on the island of Öland, south Sweden. The samples represented three generations: haploid maternal gametophytes; diploid sporophytes; and haploid sporelings. KEY RESULTS: Admixture analyses of SNP genotypes identified a majority as pure H. lutescens or H. sericeum, but 76 samples were identified as mildly admixed (17 %) and 17 samples (3.8 %) as strongly admixed. Admixed samples were represented in all three generations in several populations. Hybridization and introgression were bidirectional. CONCLUSIONS: Our results demonstrate that admixed genomes are transferred between the generations, so that the populations behave as true hybrid zones. Earlier studies of sympatric bryophyte populations with admixed individuals have not been able to show that admixed alleles are transferred beyond the first generation. The presence of true hybrid zones has strong evolutionary implications because genetic material transferred across species boundaries can be directly exposed to selection in the long-lived haploid generation of the bryophyte life cycle, and contribute to local adaptation, long-term survival and speciation.

Balance between inbreeding and outcrossing in a nannandrous species, the moss Homalothecium lutescens.

Epiphytic dwarf males on the females present a possible solution to the problem of short fertilization distances in mosses. However, leptokurtic spore dispersal makes dwarf males likely to be closely related to their host shoot, with an accompanying risk of inbreeding. The capacity of a female to harbour a high number of different dwarf males suggests that there may be mechanisms in place that counteract inbreeding, such as polyandry and post-fertilization selection. We have genotyped sporophytes, female host shoots and dwarf males in four populations of the moss Homalothecium lutescens. We found no evidence of selective sporophyte abortion based on level of heterozygosity. The occurrence of entirely homozygous sporophytes together with significantly positive inbreeding coefficients in three of the populations (mean FIS between 0.48 and 0.64) suggest frequent mother-son mating events. However, 23% of all sampled sporophytes had a higher level of heterozygosity compared with the mean expected heterozygosity at the population level. Polyandry was frequent, on average 59% of the sporophytes on a female shoot were sired by distinct fathers. In conclusion, sporadic fertilizations by dwarf males originating from nonhost female shoots appear to counteract strong inbreeding.

Maternal transmission of cytoplasmic DNA in interspecific hybrids of peat mosses, Sphagnum (Bryophyta).

The progeny of spontaneous interspecific hybrid sporophytes of Sphagnum were used to analyse the inheritance of cytoplasmic DNA. The analysis showed that only the female parent donated chloroplasts and mitochondria in Sphagnum hybrids. Thus, this is the first study demonstrating maternal cytoplasmic inheritance in a nonvascular land plant. This finding has important implications for phylogenetic reconstructions utilizing chloroplast and mitochondrial DNA sequences as well as for the evolution of cytoplasmic inheritance in relation to the life cycle of land plants.

Genetic diversity of the epiphytic bryophyte Leucodon sciuroides in formerly glaciated versus nonglaciated parts of Europe.

Twelve populations of the epiphytic bryophyte Leucodon sciuroides from three major regions representing formerly glaciated and nonglaciated regions of Europe were screened for polymorphisms at 15 putative isozyme loci. The populations clustered into three distinct groups consisting of: (i) a single population from Crete, representing a cryptic unknown taxon; (ii) four Scandinavian populations and two populations from northern Greece; and (iii) the remaining populations from mainland Greece and Crete. The Scandinavian populations were genetically depleted compared with most Greek populations, thus fitting the expectation of generally lower levels of variation in formerly glaciated areas. The transition zone between genetically diverse and depleted populations appears to be located through northern Greece, coinciding with the northern limit of the Mediterranean region. This indicates that genetic variation was lost in populations at the northern limit of glacial refugia. The two groups of populations fit a progenitor-derivative model. They also have contrasting reproductive strategies: the Mediterranean populations reproduce sexually, whereas the other populations propagate vegetatively. Epiphytic species, growing on substrates that are limited in space and time, appear to be especially vulnerable to loss of genetic variation. Lack of genetic variation and therefore low adaptability to increased levels of atmospheric pollution may explain why many epiphytic lichen and bryophytes, including L. sciuroides, are declining over much of Europe.