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Transcript
Mechanisms of Plant Evolution Special topic: the origin of Angiosperms Special topic: origin of Angiosperms Two concurrent models for the origin of Angiosperms Paleoherb Hypothesis: primitive angiosperms with short life cycle and mixed monocot- / dicot-traits as origin (Taylor und Hickey). Pro: 18S, flower morphology Woody Magnoliid Hypothese: Magnolia like trees as origin (Doyle und Donoghue). Pro: non-fused petals, no sepals, whorls of flower organs, living fossiles with few species Special topic: origin of Angiosperms •Missing links: Fossile records for the transition between Gymnoand Angiosperms are lacking. How did the first flower look like? •Gnetales: Primitive gymnosperms that are almost extinct, but display at the same time typical Angiosperm traits: Archaefructus. In 1998 the so far oldest Angiosperm fossile was discovered. Archaefructus (Clusiaceae) – Sun et al. In the Search of the First Flower, Science 282, 1692ff. This plant was very close to the Gnetales – primitive gymnosperms that are almost extinct. •Angiosperm traits in Gnetales: Double fertilization, reticulate leaves, flower-like sporangiophores, xylem with trachea. Special topic: origin of Angiosperms •Gnetales - advanced: reticulate leaves, double fertilization, „flowers“, xylem with trachea. •Gnetales - primitive: seeds not surrounded by maternal tissue (gymnosperms), anthers leaf like, no separation theca-filament, carpels open, not fused into a node. Gnetum gnemon. Male and female flowers of Gnetum gnemon –Note the reticulate leaves that are typical for Angiosperms. Evolutionary series for anthers and carpels (primitive left, advanced right). Gnetum is a gymnosperm with primitive anthers and carpels. By the way: this primordial plant exists in Germany only in the Botanical Garden of Karlsruhe! •Missing link: Gnetum gnemon combines advanced and primitive traits. It might therefore be near the branching point between gymno- and angiosperms. Special topic: origin of Angiosperms •Archaefructus: The flower (e.g. fused anthers and carpels) is much more advanced than that of Gnetum, but the morphology places Archaefructus in the Gnetales. •Gnetales: By Gnetum falls gymnosperms. Archaefructus. According to molecular traits, the Gnetales are clearly in the gymnosperms (right), according to morphological traits, they are close to the angiosperms. Archaefructus is closer than Gnetum gnemon – molecular data do not exist in fossiles! molecular phylogeny, clearly into the •Archaefructus is the oldest known angiosperm: Flower organs partially more modern than Magnoliidae, other traits are more primitive. 1:0 for Woody Magnoliid! Special topic: origin of Angiosperms OLD conifers club dicots mosses paleoherbs ferns cycads Magnoliids monocots Shocker: Monocots First loss of cambium, herbs cambium •Monocot-first hypothesis: no common ancestor for gymno- and angiosperms. Semiaquatic club mosses yield monocots, thus no selection for cambium, cambium was invented a second time giving rise to the rapid development of the dicots NEW conifers club dicots mosses ferns Magnoliids paleoherbs cycads monocots •Traditional: seed plants monophyletic, cambium in common ancestor of gymnoand angiosperms, tree-habit basal. Later lost in monocots and paleoherbs. Alisma (monocot) •support: genetics, embryology Isoetes / monocots, dicot cotyledons not homologous to „monocot“, primitive monocots amphibic, linear leaf venation from pin-formed telomes, primitive order of vasculature in monocots cambium twice Isoetes (club moss)