Origin of Angiosperms

Geologic time scale

Pollen evidence: 126 mya (Lower Cretaceous Period) Macrofossil evidence (Archaeofructus): 130 mya (Lower Cretaceous)

WHERE: West Gondwana (South

America and Africa)

Archaefructus liaoningensis

Pioneer plants in the Cretaceous: figs, planes (Platanus), and magnolias

Possible reasons for rapid radiation of angiosperms:

Co-evolution with many insect groups Shorter life cycle Diverse habitat types and edaphic factors

Angiosperm diversity
at least 260,000 living species

classified in 453 families (Judd et al., 2002; APG II, 2003; Soltis et al., 2005) epiphytes, floating and rooted aquatics, terrestrial (herbs, shrubs, vines, lianas, giant trees), parasitic forms Variation in size, longevity & overall form

Angiosperm synapomorphies (derived,

shared characters):
1) Ovules enclosed within carpels
2) Double fertilization 3) Stamens with two pairs of pollen sacs 4) Features of gametophyte structure and development 5) Phloem tissue (sieve tubes and companion cells)

IsoetesMonocotyledon theory (Campbell, 1928)

Similarities in: habitat (humid tropics); habit; embryo & anatomy of older sporophyte

Gnetalean ancestry
(Wettstein, 1901) Similarities: two cotyledons, vessels in 2ndary wood, two integuments, net-veined leaves

Coniferalian ancestry of Amentiferae (Eichler, 1875;

Engler & Prantl, 1924)
Similarity in: the inflorescence (simple & naked flowers) of amentifers (Casuarinaceae, Salicaceae, Fagaceae) and the conifers; seeds of Juniperus resembles that of Amentifers.

Pteridospermean ancestry (Long, 1966;

Cronquist)
Reticulate venation; monopodial branching; presence of cambium; microsporophylls & megasporophylls on the same plant; stelar structure; absence of vessels in certain angiosperms, like Winteraceae & Nymphaeaceae.

Caytonialean ancestry
(Thomas, 1925; Stebbins, 1974)

Caytoniales- fossil plants in the Middle Jurassic; had angiosperm-like anthers similar to branched stamens in Ricinus & Hypericum; ovules enclosed by cupule; netlike venation

Hypericum stamen

Caytonia. (A) Sagenopteris foliage; (B) Caytonanthus microsporophyll with synangia; (C) Caytonia megasporophyll and longitudinal section of cupule showing ovule position and orientation.

The Woody Magnoliid Hypothesis -- (Doyle and

Donoghue) That the earliest angiosperms had similar morphology to living members of Laurales & Magnoliales; LONGHELD PARADIGM; supported by Takhtajan, Hutchinson, Bessey, and Stebbins; also known as Euanthial theory

The strobilus-like flower and primitive carpels of Magnolia.

The Paleoherb Hypothesis

(Taylor and Hickey, paleobotanists) That the basal angiosperms are tropical paleoherbs, a group of flowering plants with uncomplicated flowers and a mix of monocot and dicot features, & rapid life cycle.

PALEOHERB HYPOTHESIS

Aristolochiales

Piperales

Two competing hypotheses

Amborella hypothesis
Endemic to cloud forest of New Caledonia
described in the mid-19th century by Baillon (1869); classified as one of the basal angiosperms, together w/ Laurales (Cronquist, 1981) Has spirally-arranged floral organs Carpels closed only by secretion rather than by fused tissue, an ancestral feature (i.e., plesiomorphy) Without vessels

Amborella trichopoda

Amborella

Austrobaileya

Sarcandra chloranthoides

Austrobaileya

Leaf-fish stamen

Ceratophyllum hypothesis (Chase et al.,1993

Ceratophyllum as sister to all other angiosperms based on molecular data (rbcL gene) Submerged plants, flowers small and dioecious
Pistillate flower

Tricolpate pollen

Monosulcate pollen

Examples of primitive angiosperms

Englerian school:
Simple plants are primitive (Amentiferae) & complex ones are advanced Based on progressive evolution

Ranalian school:
Ranales group is primitive Evolution proceeded in progressive & retrogressive manner

Winteraceae(Cronquist, 1988;

Thorne, 1983): Drimys, Bubbia, Exospermum- vesselless, conduplicate carpel

Magnoliaceae (Hutchinson,

1959)- elongated floral axis, spirallyarranged stamens & carpels; laminar stamen; very small embryo & abundant endosperm

Magnolia purpurea

Ranales: Ranunculus

Ranales: Berberidaceae