The life form spectra of communities

We pointed out earlier the crucial importance of geographic isolation in allowing populations to diverge under selection. The geographic distributions of species, genera, families and even higher taxonomic categories of plants and animals often reflect this geographic divergence. All species of lemurs, for example, are found on the island of Madagascar and nowhere else. Similarly, desert tropical rainforest aquatic biomes?

230 species in the genus Eucalyptus (gum tree) occur naturally in Australia (and two or three in Indonesia and Malaysia). The lemurs and the gum trees occur where they do because they evolved there - not because these are the only places where they could survive and prosper. Indeed, many Eucalyptus species grow with great success and spread rapidly when they have been introduced to California or Kenya. A map of the natural world distribution of lemurs tells us quite a lot about the evolutionary history of this group. But as far as its relationship with a biome is concerned, the most we can say is that lemurs happen to be one of the constituents of the tropical rainforest biome in Madagascar.

Similarly, particular biomes in Australia include certain marsupial mammals, while the same biomes in other parts of the world are home to their placental counterparts. A map of biomes, then, is not usually a map of the distribution of species. Instead, we recognize different biomes and different types of aquatic community from the types of organisms that live in them. How can we describe their similarities so that we can classify, compare and map them? In addressing this question, the Danish biogeographer Raunkiaer developed, in 1934, his idea of 'life forms', a deep insight into the ecological significance of plant forms (Figure 1.19). He then used the spectrum of life forms present in different types of vegetation as a means of describing their ecological character.

Plants grow by developing new shoots from the buds that lie at the apices (tips) of existing shoots and in the leaf axils. Within the buds, the meris-tematic cells are the most sensitive part of the whole shoot - the 'Achilles' heel' of plants. Raunkiaer argued that the ways in which these buds are protected in different plants are powerful indicators of the hazards in their environments and may be used to define the different plant forms (Figure 1.19). Thus, trees expose their buds high in the air, fully exposed to the wind, cold and drought; Raunkiaer called them phanerophytes (Greek phanero, 'visible'; phyte, 'plant'). By contrast, many perennial herbs form cushions or tussocks in which buds are borne above ground but are protected from drought and cold in the dense mass of old leaves and shoots (chamaephytes: 'on the ground plants'). Buds are even better protected when they are formed at or in the soil surface (hemicryptophytes: 'half hidden plants') or on buried dormant storage organs (bulbs, corms and rhizomes -cryptophytes: 'hidden plants'; orgeophytes: 'earth plants'). These allow the plants to make rapid growth and to flower before they die back to a dormant state. A final major category consists of annual plants that depend wholly on dormant seeds to carry their populations through seasons of drought and cold (therophytes: 'summer plants'). Therophytes are the plants of deserts (they make up nearly 50% of the flora of Death Valley, USA), sand dunes and repeatedly disturbed habitats. They also include the annual weeds of arable lands, gardens and urban wastelands.

But there is, of course, no vegetation that consists entirely of one growth form. All vegetation contains a mixture, a spectrum, of Raunkiaer's life forms. The composition of the spectrum in any particular habitat is as good a shorthand description of its vegetation as ecologists have yet managed to devise. Raunkiaer compared these with a 'global spectrum' obtained by sampling from a compendium of all species known and described in his time (the Index Kewensis), biased by the fact that the tropics were, and still are, relatively unexplored. Thus, for example, we recognize a chaparral type of vegetation when we see it in Chile, Australia, California or Crete because the life form spectrums are similar. Their detailed taxonomies would only emphasize how different they are.

Faunas are bound to be closely tied to floras - if only because most herbivores are choosy about their diet. Terrestrial carnivores range more widely than their herbivore prey, but the distribution of herbivores still gives the carnivores a broad vegetational allegiance. Plant scientists have tended to be keener on classifying floras than animal scientists on classifying faunas, but one interesting attempt to classify faunas compared the mammals of forests in Malaya, Panama, Australia and Zaire (Andrews et al., 1979). They were classified into carnivores, herbivores, insectivores and mixed feeders, and these categories were subdivided into those that were aerial (mainly bats and flying foxes), arboreal (tree dwellers), scansorial (climbers) or small ground mammals (Figure 1.20). The comparison reveals some strong contrasts and similarities. For example, the ecological diversity spectra for the Australian and Malayan forests were very similar despite the fact that their faunas are taxonomically very distinct - the Australian mammals are marsupials and the Malaysian mammals are placentals.

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