title: Escaping the lianoid habit: evolution of shrub-like growth forms in Aristolochia subgenus Isotrema (Aristolochiaceae).
abstract: OBJECTIVE
  A large range of growth forms is a notable aspect of angiosperm diversity and arguably a key element of their success. However, few studies within a phylogenetic context have explored how anatomical, developmental, and biomechanical traits are linked with growth form evolution. Aristolochia (∼500 species) consists predominantly of climbers, but a handful of shrub-like species are known from Aristolochia subgenus Isotrema (hereafter, shortened to Isotrema). We test hypotheses proposing that the establishment of functional traits linked to lianescence might limit the ability to evolve structurally diverse growth forms, particularly self-supporting forms. •
  
  
  METHODS
  We focus on the origin of the shrub habit in Isotrema, from which we sampled representatives from climbing to self-supporting forms. Morphological, anatomical, and biomechanical characters are optimized on a chloroplast- and nuclear-derived phylogeny. •
  
  
  RESULTS
  Character-state reconstructions revealed that the climbing habit is plesiomorphic in Isotrema and shrub-like forms are derived from climbers. However, shrubs do not constitute a monophyletic group. Both shrubs and climbers show large multiseriate rays, but differ in terms of vessel size and proportion of fibers and soft tissues. •
  
  
  CONCLUSIONS
  We suggest that while shrub-like species might have partly escaped from the constraints of life as lianas; their height size and stability are not typical of self-supporting shrubs and trees. Shrubs retained lianoid stem characters that are known to promote flexibility such as ray parenchyma. The transitions to a shrub-like form likely involved relatively simple, developmental changes that may be attributed to heterochronic processes.
authors.affiliation: Institut für Botanik, Technische Universität Dresden, D-01062 Dresden, Germany. sarah.wagner@ecemp.tu-dresden.de
authors.forename: Sarah T
authors.initials: ST
authors.lastname: Wagner
authors.forename: Sandrine
authors.initials: S
authors.lastname: Isnard
authors.forename: Nick P
authors.initials: NP
authors.lastname: Rowe
authors.forename: Marie-Stéphanie
authors.initials: MS
authors.lastname: Samain
authors.forename: Christoph
authors.initials: C
authors.lastname: Neinhuis
authors.forename: Stefan
authors.initials: S
authors.lastname: Wanke
mesh_terms: D030022:Aristolochia / Q000033:anatomy & histology* / Q000254:growth & development* / Q000502:physiology; D005075:Biological Evolution; D001696:Biomechanical Phenomena; D030541:Databases, Genetic; D017753:Ecosystem; D055119:Elastic Modulus; D010802:Phylogeny; D018547:Plant Stems / Q000502:physiology; D052584:Xylem / Q000033:anatomy & histology