Angiosperm origins and relationships of major lineages

Vessels and tracheids

Companion cells

Pollen columellae

Carpel and integuments

Fertilisation

Development of Fruits

Cycads

Conifers

Ginkgos

Gnetaceae

Bennettitales

Archaefructus

The oldest angiosperm

Early lineages (i)

Early lineages (ii) Chloranthaceae Piperaceae

Earliest Angiosperms Perennial Evergreen Woody Vascular bundles in a ring, Vessel-less wood Alternate, simple, coriaceous leaves with entire or sparsely-toothed margins Lacking stipules Lacking ethereal oils

Earliest Flowers Insect-pollinated Radially symmetrical Several - many tepals, spirally-arranged, free and distinct Bisexual Stamens poorly differentiated into anther and filament Anthers with four microsporangia opening by longitudinal slits Superior ovaries Pollen monosulcate and often lacking well-developed columellae Carpels fused by secretion, with an elongated stigma and a style poorly differentiated from ovary. Placentation marginal

Dicotyledons Monocotyledons Embryo with 2 cotyledons Primary root persistent Herbs, shrubs or trees Stele cylindrical Pollen tricolpate Monocotyledons Embryo with 1 cotyledon Primary root replaced by adventitious roots Mostly herbs Vascular bundles scattered Pollen monocolpate

Monocolpate and Tricolpate

Anomalous Dicots Flowers in 3’s or with numerous parts Poorly differentiated anthers Poorly developed styles and elongate stigmas Pollen monocolpate or inaperturate Ethereal oils in vegetative parts

Magnoliaceae

Lauraceae

Piperaceae

Nymphaceae

A summary of the major groups that occurred in greater than 50% of the 1000 jackknife replicates (from Soltis et al. 2000).

ANITA Amborellaceae Nymphaceae Illiciaceae Trimeniaceae Austrobaileyaceae

MAGNOLIIDS Chloranthaceae Magnoliaceae Annonaceae Myristicaceae Lauraceae Piperaceae Aristolochiaceae Ceratophyllaceae

Amborellaceae New Caledonia

Amborella trichopoda

Amborella trichopoda Shrubby Lacks vessels No ethereal oils Radial flowers with 7-11 tepals Dioecious Ascidiate carpels

Nymphaceae Victoria amazonica

Nymphaceae

Austrobaileyales llliciaceae

Austrobaileyales Trimeniaceae

Austrobaileyales Austrobaileyaceae

Chloranthaceae

Piperaceae

Aristolochiaceae

Lauraceae

Magnoliaceae

Annonaceae

Myrtisticaceae

Ceratophyllaceae

Basal Angiosperms Monocots Eudicots Basal Eudicots Rosids Asterids

Monocot autapomorphies Embryo with 1 cotyledon Vascular bundles scattered Primary root replaced by adventitious roots Leaves with parallel venation and sheathing bases Herbaceous habit DNA sequences

Basal Angiosperms Monocots Eudicots Basal Eudicots Rosids Asterids

Eudicot autapomorphies Tricolpate pollen Sequences from rbcL, atpB, 18S rDNA Floral parts in whorls Slender filaments

Basal Angiosperms Monocots Eudicots Basal Eudicots Rosids Asterids

Core Eudicots autapomorphies Sequences from rbcL, atpB, 18S 4- or 5-merous flowers with calyx and corolla Placentation usually axile

Basal Angiosperms Monocots Eudicots Basal Eudicots Rosids Asterids

Eurosids autapomorphies Sequences from rbcL, atpB, matK, 18S Nitrogen fixers in clade of Fabales, Rosales, Cucurbitales and Fagales

Basal Angiosperms Monocots Eudicots Basal Eudicots Rosids Asterids

Asterids autapomorphies Sequences from rbcL, atpB, matK, ndhF 18S Ovules with a single integument

Basal Angiosperms Monocots Eudicots Basal Eudicots Rosids Asterids

Core Asterids autapomorphies Sympetalous corolla Number of petals = number of stamens Epipetalous stamens Gynoecium of 2 fused carpels DNA sequences