The first flowering plants 1. 2. 3. 4. 5. The most primitive living angiosperms The shared primitive characters Fossils of the most primitive angiosperms Living sister groups Extinct sister groups General Angiosperm Relations --- Soltis et al. 2008 Amborella Amborella - flowers Amborella Amborella Nymphaea Nymphaea Nymphaea Hydatella Austrobaileya blooming liana at UVM Austrobaileya flowers native to Queensland, northeastern Australia The first flowering plants 1. 2. 3. 4. 5. The most primitive living angiosperms The shared primitive characters Fossils of the most primitive angiosperms Living sister groups Extinct sister groups Inferred ancestral features of angiosperms (from living groups) 1) 2) 3) 4) 5) 6) more than two whorls (or series) of tepals and stamens stamens with protruding adaxial or lateral pollen sacs several free, ascidiate carpels closed by secretion extended stigma extragynoecial compitum one or several ventral pendent ovule(s) equivocal: 1) bisexual vs. unisexual fl owers 2) whorled vs. helical attachment to receptacle 3) presence vs. absence of tepal differentiation 4) anatropous vs. orthotropous ovules. Simple flowers of the basal groups are reduced rather than primitively simple. Endress and Doyle 2009 Distribution of ascidiate carpels Trimenia has ascidiate carpels. And the pistils of Amborella are taken as simple, ascidiate carpels. The gynoecium of Myristica arises as a single ascidiate carpel, then develops a cleft. Endress’s sequence of evolution of the early angiosperm carpel. Secretion in blue, post-genital fusion in red. Amborella Amborellaceae Austrobaileya Austrobaileyaceae Asimina Annonaceae Nymphaea Nymphaeaceae Myristica Myristicaceae So what the heck is an extragynoecial compitum? Major changes in flower morphology (and most primitive states) The first flowering plants 1. 2. 3. 4. 5. The most primitive living angiosperms The shared primitive characters Fossils of the most primitive angiosperms Living sister groups Extinct sister groups The Cretaceous Archaefructus – early Aptian (124my) Nymphaeales First angiosperm fossils --- in phylogenetic context from Doyle and Endress, 2010 First angiosperm fossils --- sequence of diversification from Doyle and Endress, 2010 The first flowering plants 1. 2. 3. 4. 5. The most primitive living angiosperms The shared primitive characters Fossils of the most primitive angiosperms Living sister groups Extinct sister groups Gnetum The first flowering plants 1. 2. 3. 4. 5. The most primitive living angiosperms The shared primitive characters Fossils of the most primitive angiosperms Living sister groups Extinct sister groups Glossopteris Pentaxylon Bennetitales: Cycadeoidea Bennetitales: Williamsonia Caytonia So, Bailey’s plicate (conduplicate) carpel may be a valid inference based on the Magnoliid carpel, but derived from a plicate carpel. Bailey’s drawings of carpels from the Winteraceae, Magnoliids The first flowering plants 1. 2. 3. 4. 5. The most primitive living angiosperms The shared primitive characters Fossils of the most primitive angiosperms Living sister groups Extinct sister groups Extra slides Peltaspermum Callistophyton FLOWER FORM AND MOLECULAR DEVELOPMENT (A) Classic ABCE model (A)Evolution of MADS genes (I) ABC model developed for core eudicots (II)“shifting boundary model” applied to some basal eudicots and monocots (III) “fading borders”model proposed for basal angiosperms Ascidiate carpels with an extragynoecial compitum….. compitum: a tract of transmission tissue in the gynoecium that is common to all the carpels of the one flower and that allows pollen landing on any one stigma or part of a stigma to fertilise ovules in any carpel Plicate carpels.