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5.3 Class Magnoliopsida – flowering plants - Cambridge University ...

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Insect pollination is closely associated with the origin and subsequent<br />

diversification of flowers. However, it is important to remember<br />

that insect pollination is associated with several other groups of seed<br />

<strong>plants</strong>, both living and extinct: Bennettitales, Gnetales, Cheirolepidaceae<br />

(extinct conifers), Cycadales and Medullosales (seed ferns).<br />

Insects grew in diversity with the origin of seed <strong>plants</strong> in the Late<br />

Devonian and this increase of diversity with the origin of flowers<br />

is just part of a continuous trend. Indeed there is some evidence<br />

to indicate a temporary decline in insect diversity as <strong>flowering</strong> <strong>plants</strong><br />

became more abundant in the Cretaceous. Nevertheless, flowers diversified<br />

in parallel with particular groups of pollinator specialists,<br />

the bees (Apoidea/Apidae), the pollen wasps (Vespidae: Masarinae),<br />

brachyceran flies (Acroceridae, Apioceridae, Bombyliidae, Empididae,<br />

Nemestrinidae, Stratiomyidae and Syrphidae) and the moths and butterflies<br />

(Lepidoptera). The evolution of a bisexual reproductive axis<br />

was a crucial event.<br />

Several trends in floral evolution can be discerned. Primitive flowerseitherlackaperianthorhaveoneinwhichthereisasingle<br />

whorl of tepals. In the perianth there has been a trend from having<br />

a perianth in which distinct whorls are not clearly differentiated to<br />

clear specialisation of a distinct calyx and corolla. The calyx protects<br />

against drought, temperature shock and predatory insects and the<br />

corolla attracts and controls pollinators. Both calyx and corolla may<br />

have been derived from tepals, but it is likely that in some groups,<br />

such as the buttercups, the petals originated from stamens, to which<br />

they are anatomically similar. In the peony, Paeonia, there is a gradual<br />

transition from leaves, through modified leaves on the <strong>flowering</strong> stem<br />

called bracts, into the perianth with parts at first sepal-like and then<br />

petal-like. Generally there has been a trend for the greater integration<br />

of the floral parts with greater precision in number and placement as<br />

flowers have become specialised to particular patterns of pollination.<br />

An important aspect of the diversification of <strong>flowering</strong> <strong>plants</strong> and<br />

their evolutionary success has been their vegetative flexibility. They<br />

have evolved into a bewildering range of forms through the activity<br />

of sub-apical and intercalary meristems. Flowering <strong>plants</strong> also have<br />

a greater capacity for elongation of cells, including root hairs and<br />

trichomes. One of the most important vegetative specialisations has<br />

been their possession of vessels in their wood, permitting more efficient<br />

water transport and hence greater photosynthetic rates, fast<br />

growth rates and earlier maturation. There have been significant<br />

adaptations in seasonal habitats; for example, although a few non<strong>flowering</strong><br />

<strong>plants</strong> are deciduous this has been a particular <strong>flowering</strong>plant<br />

trait that has adapted them to high latitudes or seasonally-dry<br />

environments.<br />

One aspect to the burgeoning biotic diversity of <strong>flowering</strong> <strong>plants</strong><br />

from the Cretaceous onwards was their great expansion of chemical<br />

diversity. New ranges of what have been called secondary compounds<br />

provided attractants for pollinators or seed dispersers, and repellants<br />

and toxic compounds to inhibit herbivores.<br />

<strong>5.3</strong> CLASS MAGNOLIOPSIDA <strong>–</strong> FLOWERING PLANTS 225<br />

(a)<br />

(b)<br />

Figure 5.55. Specialisation in<br />

the perianth in the Asparagales:<br />

(a) Belamcanda; (b) Kniphofia.

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