Nomenclature and typification in Verbascum (Scrophulariaceae) from North Africa.

The progress of taxonomic work on native Verbascum L. taxa found in Morocco led to a search for reference specimens in various herbaria. This process was extended to the taxa found in the other four countries of North Africa (Algeria, Tunisia, Libya, and Egypt), which make up the southern shore of the Mediterranean basin. Numerous names were identified as needing typification or requiring corrections of their earlier lectotypifications in order to stabilize their nomenclature and provide a better definition of each taxon. As a result, lectotypes are now designated for 35 names, a neotype is proposed for V.ballii (Batt.) Hub.-Mor., and second-step lectotypes are proposed for V.faureisubsp.acanthifolium (Pau) Benedí & J.M.Monts. and V.pinnatisectum (Batt.) Benedí. Comments have been added for each typified name. Known isolectotypes are also mentioned whenever possible. Furthermore, some new combinations are proposed in this paper, namely V.longirostrevar.antiatlantica (Emb.) Khamar, comb. nov., V.longirostrevar.atlantica (Maire) Khamar, comb. nov., and V.longirostrevar.hoggarica (Maire) Khamar, comb. nov.

The diversity and evolution of pollination systems in large plant clades: Apocynaceae as a case study.

Background and Aims: Large clades of angiosperms are often characterized by diverse interactions with pollinators, but how these pollination systems are structured phylogenetically and biogeographically is still uncertain for most families. Apocynaceae is a clade of >5300 species with a worldwide distribution. A database representing >10 % of species in the family was used to explore the diversity of pollinators and evolutionary shifts in pollination systems across major clades and regions. Methods: The database was compiled from published and unpublished reports. Plants were categorized into broad pollination systems and then subdivided to include bimodal systems. These were mapped against the five major divisions of the family, and against the smaller clades. Finally, pollination systems were mapped onto a phylogenetic reconstruction that included those species for which sequence data are available, and transition rates between pollination systems were calculated. Key Results: Most Apocynaceae are insect pollinated with few records of bird pollination. Almost three-quarters of species are pollinated by a single higher taxon (e.g. flies or moths); 7 % have bimodal pollination systems, whilst the remaining approx. 20 % are insect generalists. The less phenotypically specialized flowers of the Rauvolfioids are pollinated by a more restricted set of pollinators than are more complex flowers within the Apocynoids + Periplocoideae + Secamonoideae + Asclepiadoideae (APSA) clade. Certain combinations of bimodal pollination systems are more common than others. Some pollination systems are missing from particular regions, whilst others are over-represented. Conclusions: Within Apocynaceae, interactions with pollinators are highly structured both phylogenetically and biogeographically. Variation in transition rates between pollination systems suggest constraints on their evolution, whereas regional differences point to environmental effects such as filtering of certain pollinators from habitats. This is the most extensive analysis of its type so far attempted and gives important insights into the diversity and evolution of pollination systems in large clades.

Stable epigenetic effects impact adaptation in allopolyploid orchids (Dactylorhiza: Orchidaceae).

Epigenetic information includes heritable signals that modulate gene expression but are not encoded in the primary nucleotide sequence. We have studied natural epigenetic variation in three allotetraploid sibling orchid species (Dactylorhiza majalis s.str, D. traunsteineri s.l., and D. ebudensis) that differ radically in geography/ecology. The epigenetic variation released by genome doubling has been restructured in species-specific patterns that reflect their recent evolutionary history and have an impact on their ecology and evolution, hundreds of generations after their formation. Using two contrasting approaches that yielded largely congruent results, epigenome scans pinpointed epiloci under divergent selection that correlate with eco-environmental variables, mainly related to water availability and temperature. The stable epigenetic divergence in this group is largely responsible for persistent ecological differences, which then set the stage for species-specific genetic patterns to accumulate in response to further selection and/or drift. Our results strongly suggest a need to expand our current evolutionary framework to encompass a complementary epigenetic dimension when seeking to understand population processes that drive phenotypic evolution and adaptation.

Evolution of shrub-like growth forms in the lianoid subfamily Secamonoideae (Apocynaceae s.l.) of Madagascar: phylogeny, biomechanics, and development.

Lianas are common in the Apocynaceae s.l. and are predominant in the subfamily Secamonoideae. Shrub-like taxa are rare within this subfamily but occur in Malagasy genera such as Secamone, Secamonopsis, and Pervillaea. We explored the evolutionary appearance of shrub-like growth forms in Malagasy Secamonoideae through a molecular phylogeny using chloroplastic sequences. The phylogeny revealed several independent appearances of shrub-like growth forms within the Secamonoideae. Biomechanics and development of the shrub-like growth form were detailed in one species, Secamone sparsiflora, which has upright and self-supporting young stems that become procumbent in older stages of development. Biomechanical investigations revealed characteristics atypical of both lianas and self-supporting shrubs. Anatomical development in S. sparsiflora is initially similar to lianas in the same clade but shows potentially neotenic retention of juvenile wood development for most of the growth trajectory. The results suggest that evolution of lianescence can carry a high degree of specialization and developmental burden that might limit evolution back to self-supporting growth forms. Under certain geographic and ecological conditions, such as geographic isolation, xeric conditions and/or reduced biotic competition, escapes from lianescence to other growth forms can occur in some angiosperm groups via relatively simple heterochronic shifts of mechanically significant growth processes.