ENDRESS1, MARY E.*, BENGT SENNBLAD 2, RAYMOND W. J. M. VAN DER HAM3, SIWERT NILSSON4, KURT POTGIETER5, LAURE CIVEYREL6, MARK CHASE7, JEFFREY JOSEPH7, MARTYN POWELL7, DAVID LORENCE8, ANTON IGERSHEIM9, AND VICTOR A. ALBERT10. 1Institute of Systematic Botany, University of Zurich, Zurich, Switzerland; 2Department of Systematic Botany, Uppsala University, Uppsala, Sweden; 3Nationaal Herbarium Nederland, Leiden, Netherlands; 4Palynological Laboratory, Swedish Museum of Natural History, Stockholm, Sweden; 5Department of Plant Biology, University of Illinois, Urbana, Illinois, U.S.A.; 6Laboratoire d'Ecologie Terrestre, Universite Paul Sabatier, Toulouse, France; 7Royal Botanic Gardens, Kew, Richmond, Surrey, United Kingdom; 8National Tropical Botanical Garden, Lawai, Hawaii, U.S.A.; 9Institute of Botany, University of Vienna, Vienna, Austria; 10Botanical Laboratory, University of Copenhagen, Copenhagen, Denmark. - Arils, wings, and other sneaky things: coming to terms with the Alyxieae (Apocynaceae).
Tribal delimitation in the Apocynaceae has traditionally been based on
fruit and seed characters. These were especially important in the
basalmost subfamily, Rauvolfioideae (usually called
"Plumerioideae"), where the flowers are often superficially
similar, with relatively unspecialized organs and few distinguishing
morphological characteristics. Results of various, mostly molecular,
studies in recent years, however, suggest that the fruit-based
classifications did not yield monophyletic groups. Here we focus on
one such group of the Rauvolfioideae, which has usually been treated
as a tribe called the "Rauvolfieae" or "Alyxieae",
and characterized by having drupaceous fruits. This study is based on
a combined analysis from five data sets, including four genes (matK,
rbcL, trnL intron and trnL-F spacer) and 54 morphological characters.
In the consensus trees from each individual data set the bulk of the
genera fall within two clades, which have been re-circumscribed as
Alyxieae and Vinceae. In the combined matrix consensus tree the
traditional "Alyxieae" is polyphyletic, with genera
dispersed over 4 separate clades. This exemplifies the ease with which
fruits may switch back and forth from dry to fleshy and/or dehiscent
to indehiscent, or seeds independently evolve similar structures to
aid in dispersal in response to selective pressures. It also bodes ill
for delimitation of higher taxonomic categories based on traditional
fruit and seed characters elsewhere in the family, and should serve as
a warning in other families in the Gentianales as well, such as
Rubiaceae, where fruit characters have played an important role in
classification. Combining morphological and molecular methods allows
us to glimpse how much more intricate the evolutionary pathways are
than previously imagined and provides a groundwork on which to study
character evolution.
Key words: Alyxieae, Apocynaceae, fruit and seed character evolution, Rauvolfieae, Rauvolfioideae