Systematic reappraisal of marsh-orchids native to Scotland

The intensively studied Eurasian orchid genus Dactylorhiza has become a model system for exploring allopolyploid evolution, yet determining the optimal circumscriptions of, and most appropriate ranks for, its constituent taxa remain highly controversial topics. Here, novel allozyme data and detailed morphometric data for 16 Scottish marsh-orchid populations are interpreted in the context of recent DNA sequencing studies. Despite being derived from the same pair of parental species, the two allopolyploid species that currently occur in Scotland can reliably be distinguished using allozymes, haplotypes, ribotypes or sequences of nuclear genes. A modest range of diverse morphological characters are shown to distinguish the two molecularly-circumscribed species, but they have in the past been obscured by equivalent levels of infraspecific variation in characters rooted in anthocyanin pigments; these characters are better employed for distinguishing infraspecific taxa. Dactylorhiza francis-drucei (formerly D. traunsteinerioides) is confirmed as being distinct from the continental D. traunsteineri/lapponica, probably originating through allopatric isolation once the continental lineage reached Britain. All Scottish populations are attributed to the comparatively small-flowered, anthocyanin-rich subsp. francis-drucei, which includes as a variety the former D. 'ebudensis'; the less anthocyanin-rich subsp. traunsteinerioides is confined to Ireland, North Wales and northern England. In contrast with D. francis-drucei, only a minority of Scottish populations of D. purpurella are attributed to the anthocyanin-rich race, var. cambrensis. This species most likely originated through an allopolyploidy event that occurred comparatively recently within the British Isles, as it contains allozyme alleles distinctive of British rather than continental D. incarnata (its diploid pollen-parent). In contrast, the rare Scottish population of D. incarnata subsp. cruenta shares with its Irish counterparts a continental genotype, and is most likely a recent arrival in Scotland through long-distance dispersal. Among all European allotetraploid dactylorchids, D. purpurella is the species that most closely resembles D. incarnata, both molecularly and morphologically.