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

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238 ORDERING THE PATHS OF DIVERSITY<br />

Figure 5.83. Trochodendron.<br />

Figure 5.84. Gunnera.<br />

Figure 5.85. Polygonaceae:<br />

Reynoutria.<br />

Figure 5.86. Amaranthaceae:<br />

Ptilotus.<br />

trochodendrales<br />

This order has only two species of evergreen trees from east and South<br />

East Asia each in its own family, Tetracentron sinense and Trochodendron<br />

aralioides (Figure 5.83).<br />

gunnerales<br />

There are only two genera in the order. Gunnera has the familiar, massive,<br />

palmate and deeply ribbed leaves. Usually grown beside water it<br />

is also a colonist of land-slips. One advantage it has is the fixed nitrogen<br />

it gets from the symbiotic blue-green bacteria (Nostoc) that live<br />

in its exposed roots and rhizomes. It produces large strobiloid inflorescences<br />

of tiny flowers, either bisexual or unisexual. Myrothamnus,<br />

from tropical Africa and Madagscar, is a resurrection plant, appearing<br />

to dry out but able to revive and start growing again when water<br />

becomes available.<br />

Core Eudicots<br />

Core eudicots have predominantly flowers with parts in fives (pentamerous)<br />

with a clear distinction between calyx and corolla. There<br />

are two main lineages, the Rosids and Asterids, three large basal lineages,<br />

the Caryophyllales, the Santalales and the Saxifragales, and a<br />

number of others of uncertain relationship like the Berberidopsidales<br />

and Vitales. These basal orders are crassinucellate (see below).<br />

caryophyllales (catchflies, stonecrops and cacti)<br />

The Caryophyllales is a large, and an interesting group of about 4%<br />

of all <strong>flowering</strong> <strong>plants</strong> that exhibits a unique set of characters (see<br />

Figure 5.86). For example it seems to lack mycorrhizae. It comprises<br />

mostly herbs, but others are lianes or twiners and shrubs. They have a<br />

peculiar pattern of secondary growth with the production of diffuse<br />

or successive cambia, which is commonly associated with succulence<br />

and CAM photosynthesis. Many families that have a high frequency<br />

of succulence have species that are either xerophytic like the cacti<br />

(Cactaceae), stonecrops (Aizoaceae) or halophytic like the sea-lavenders<br />

and thrifts (Plumbaginaceae).<br />

Most have a campylotropous ovule in which the inner integument<br />

protrudes, and a peripheral embryo surrounding a nutritive central<br />

perisperm tissue and so they were previously called the Centrospermae.<br />

Another shared character is a peculiar type of sieve-tube plastid,<br />

and their chemistry is distinct. The core caryophyllid families are<br />

Amaranthaceae, Aizoaceae, Cactaceae, Caryophyllaceae, Didiereaceae,<br />

Molluginaceae, Nyctaginaceae, and Phytolaccaceae. Most of these have<br />

a shikimic acid biosynthetic pathway as a starting point for the synthesis<br />

of nitrogen-containing benzylisoquinoline alkaloids and the<br />

betalain pigments. The latter are utilised instead of the anthocyanins<br />

used in other <strong>flowering</strong> <strong>plants</strong>.<br />

Non-core caryophyllids are the Plumbaginaceae, Polygonaceae,<br />

Tamaricaceae, and Frankeniaceae in one clade and four families<br />

of insectivorous <strong>plants</strong> in a sister clade. The former include many<br />

halophytic <strong>plants</strong>, which are also found in core caryophyllids such<br />

as the Amaranthaceae and Chenopodiaceae (Salicornia). Many have

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