molecular phylogenetics of echinopsis (cactaceae)

1348 AMERICAN JOURNAL OF BOTANY [Vol. 99
2006 ; Hipp et al., 2007 ; Timme et al., 2007 ). It has been suggested
that hybridization plays a major role in the evolution of
Cactaceae ( Rowley, 1994 ; Machado, 2008 ) and has also been
hypothesized for Echinopsis ( Friedrich, 1974 ; Font and Picca,
2001 ; Anderson, 2005 ) and related genera ( Rowley, 1994 ). Observations
in the fi eld also suggest occasional hybridization, for
example, between E. atacamensis subsp. pasacana × E. huascha
subsp. robusta , E. terscheckii × E. candicans , E. terscheckii ×
E. huascha , and E. ancistrophora × E. saltensis (B. Schlumpberger,
unpublished data). However, at least the fi rst cross in
this list involves homoploidy (BS, unpublished data), and the
relative scarcity of tetraploidy in the phylogeny ( Fig. 1 ) thus
does not necessarily refl ect a rarity of hybridization.
Conclusions — The chloroplast DNA phylogeny presented
here for 144 species and subspecies representing all major
groups of Echinopsis sensu lato as well as all genera of Trichocereeae
demonstrates the polyphyly of this genus as currently circumscribed.
The phylogeny further reveals much convergent
evolution in fl ower traits that relate to pollination modes (see
Fig. 2A, B ) and less convergent evolution in growth forms (but
see Fig. 2C, D ). There is no conspicuous role of genome doubling
in the evolution of Echinopsis sensu lato. A new generic
classifi cation of the Trichocereeae now requires fi nding morphological
characters suffi ciently conservative for distinguishing
larger groups of species. Seed morphology and growth
form, perhaps in combination, seem promising starting points.
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