title: The evolution of dwarf shrubs in alpine environments: a case study of Alchemilla in Africa.
abstract: OBJECTIVE
  Alpine and arctic environments worldwide, including high mountains, are dominated by short-stature woody plants (dwarf shrubs). This conspicuous life form asserts considerable influence on local environmental conditions above the treeline, creating its own microhabitat. This study reconstructs the evolution of dwarf shrubs in Alchemilla in the African tropical alpine environment, where they represent one of the largest clades and are among the most common and abundant plants.
  
  
  METHODS
  Different phylogenetic inference methods were used with plastid and nuclear DNA sequence markers, molecular dating (BEAST and RelTime), analyses of diversification rate shifts (MEDUSA and BAMM) and ancestral character and area reconstructions (Mesquite).
  
  
  RESULTS
  It is inferred that African Alchemilla species originated following long-distance dispersal to tropical East Africa, but that the evolution of dwarf shrubs occurred in Ethiopia and in tropical East Africa independently. Establishing a timeframe is challenging given inconsistencies in age estimates, but it seems likely that they originated in the Pleistocene, or at the earliest in the late Miocene. The adaptation to alpine-like environments in the form of dwarf shrubs has apparently not led to enhanced diversification rates. Ancestral reconstructions indicate reversals in Alchemilla from plants with a woody base to entirely herbaceous forms, a transition that is rarely reported in angiosperms.
  
  
  CONCLUSIONS
  Alchemilla is a clear example of in situ tropical alpine speciation. The dwarf shrub life form typical of African Alchemilla has evolved twice independently, further indicating its selective advantage in these harsh environments. However, it has not influenced diversification, which, although recent, was not rapid.
authors.affiliation: Institut für Spezielle Botanik und Botanischer Garten, Johannes Gutenberg Universität, Mainz, Germany and gehrke@uni-mainz.de.
authors.forename: Berit
authors.initials: B
authors.lastname: Gehrke
authors.affiliation: Institut für Spezielle Botanik und Botanischer Garten, Johannes Gutenberg Universität, Mainz, Germany and.
authors.forename: Martha
authors.initials: M
authors.lastname: Kandziora
authors.affiliation: Institut für Spezielle Botanik und Botanischer Garten, Johannes Gutenberg Universität, Mainz, Germany and Department of Biochemistry, Stellenbosch University, Stellenbosch, South Africa.
authors.forename: Michael D
authors.initials: MD
authors.lastname: Pirie
mesh_terms: D000349:Africa; D031982:Alchemilla / Q000502:physiology*; D005075:Biological Evolution; D017753:Ecosystem; D014644:Genetic Variation; D005843:Geography; D010802:Phylogeny; D017422:Sequence Analysis, DNA; D013696:Temperature
keywords: Adaptation to alpine environments; Alchemilla; Fragariinae; Rosaceae dating; Rosoideae; alpine speciation; dwarf shrubs; secondary woodiness; species evolution