Also called:
shrubland, heathland, or chaparral

The natural development of semiarid scrublands is affected by cultivation of the land and by grazing by domestic animals, a destructive form of land use that can substantially alter vegetation. Even regions that have been used for farming only for relatively short periods have been rapidly and substantially altered. Scrublands that are dependent on nutrient-poor soil can be permanently destroyed by the addition of nutrients, such as fertilizers used for agriculture or that have entered the ecosystem as pollution. The amelioration of soil conditions permits invasion by plants from other vegetation types that can grow more vigorously than the original scrubland plants.

In most regions of Australia, agriculture, including the grazing of domestic animals, has been practiced for less than 200 years. The semiarid tropical scrublands are reasonably intact across large areas, but the more southerly chenopod scrublands have been altered markedly during the past 150 years by intense sheep grazing. Ninety percent of these valuable rangelands have degenerated to some extent, and 25 percent have been severely affected. During this process, 22 plant species have become so rare that they are considered to be in danger of extinction within the next 10 to 20 years if land-use practices are not changed. The number of palatable species seems to be declining, especially near watering points, although by contrast some unpalatable shrub species have become more common. This effect has greatly reduced the value of the vegetation as rangeland.

Large areas of mallee have been cleared to grow wheat, although the climate is so marginal that the crop often fails completely. Other Australian scrublands also continue to be cleared and fertilized for various agricultural purposes.

Broad-scale farming activity may create as well as destroy scrublands. This has happened in large areas of semiarid eastern Australia where grasslands were formerly dominant. Previously, when thick grass grew after a rain, it became fuel for wildfires, a process that favoured regrowth of the grass after the next rain but killed many shrub seedlings. After introduction of large numbers of domestic stock and eviction of the area’s Aboriginal inhabitants who had maintained the fire regime, the grass was mostly eaten by sheep and fires became less frequent. As a consequence, very large areas were invaded by “woody weeds”—unpalatable native shrubs that have created new areas of scrublands and greatly diminished the area’s grazing value.

Wherever climatic conditions are marginal for tree growth, vegetation is vulnerable to alteration by human activities. Trees disappear as a result, often to be replaced by shrubs. This vegetational degradation may have occurred so long ago as to be effectively permanent, leading modern observers to believe initially that these scrublands represent the natural, climate-induced vegetation of those areas. Scrublands of the Mediterranean region that have been affected by human activity for centuries, called by local names such as garigue, maquis, and macchia, are examples. Many areas have been converted from forest to scrubland by tree clearance, heavy grazing especially by goats, frequent burning, and consequent soil erosion. In these ways, evergreen forests of pine and oak have been replaced by scrublands in places throughout the northern Mediterranean fringe from Spain to Lebanon. Similar changes can be observed at similar latitudes as far eastward as China.

Chutes d'Ekom - a waterfall on the Nkam river in the rainforest near Melong, in the western highlands of Cameroon in Africa.
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Ecosystems

Occasionally, scrublands may result from the introduction of, and subsequent dominance by, an alien shrub species in a region in which woody plants are absent or rare. This has occurred in the grass-dominated, seasonally flooded areas of northernmost Australia to which the giant sensitive plant (Mimosa pigra) was introduced from Mexico; it since has multiplied to form extensive areas of dense, unnatural scrubland. A similar process is occurring in the Mitchell grasslands in inland northeastern Australia, an area being invaded by the introduced African shrub Acacia nilotica.

Ecological importance of fire to scrubland communities

Fire is essential to the health of most scrublands. Without periodic burning, many scrublands would alter in composition; some would gradually develop into tree-dominated vegetation. Fires serve to kill young trees or to keep them in a shrub form. They also permit reproduction by the fire-adapted scrubland flora. Most scrubland plants are well adapted to survive fire, generally in one of two ways: Some resprout from the stem base or from underground organs after the aboveground parts have been destroyed; others regenerate from seeds that have been shed on the burned ground or that lie dormant in the soil. Many scrubland plants produce seeds that can remain dormant for long periods, a useful adaptation in an environment in which periods suitable for the establishment of seedlings are infrequent and unpredictable.

Many scrubland shrubs develop a swollen stem base or an underground woody organ called a lignotuber, which, because it is at or just below the soil surface, is protected from damage even during fires that completely remove the shrub’s crown. After a fire, buds under the woody outer layers of this organ quickly grow, using the undamaged original root system; a new crown develops within a few months. Similarly, many herbaceous plants have fleshy bulbs, rhizomes, or other types of underground stem from which green shoots rapidly develop in the wake of a fire that destroyed earlier shoots. Several species are stimulated to flower by fire, so that all individuals bloom simultaneously a few weeks after being burned. An example of this behaviour is the conspicuous Australian grass trees (Xanthorrhoea).

Other plants hold their seeds in woody fruits that remain closed until being opened by fire. Only then do the seeds fall to the ground, which has been temporarily fertilized by ash and cleared of dead leaf litter, competing plants, seed-harvesting ants, and parasitic molds. Such plants typically depend on fire to provide the conditions necessary for reproduction and are unable to regenerate without appropriate intervals of burning. For example, a common and attractive shrub of coastal scrublands in eastern Australia, Banksia ericifolia, is eliminated not only if an area is burned more often than every fifth year—the time taken for seedlings to set their first seed—but also if it is burned less often than every 40 years—the plant’s life span.

Many scrubland shrubs shed seeds as they ripen, even in years when fire does not occur. The seeds remain dormant in the soil until stimulated to germinate by heat or, in some cases, by chemicals in the smoke released by a fire. Heat stimulation is required for plant germination in several families, including the bean family (many species of Acacia) and the rose family (the Californian chamiso Adenostoma fasciculatum, which is stimulated to germinate by exposure to an air temperature of 90° to 95° C [194° to 203° F]).

Biological productivity

Scrublands typically grow under conditions of high environmental stress. The typical climatic environment experienced by scrublands includes long periods of hot, dry weather in which lack of moisture is a limiting factor for plant growth. Furthermore, soil nutrient levels typically are very low. These factors restrict rates of plant photosynthesis. There may be a burst of growth during briefly favourable conditions, such as in spring in Mediterranean climatic regions, but productivity overall is not high. (For a full discussion of productivity see biosphere: The organism and the environment: Resources of the biosphere.)

The dominant shrubs may be deciduous, losing their leaves for significant periods of time when moisture is in short supply; if they are evergreen their leaves are small and tough and total leaf area is relatively low. They also typically have a substantial proportion of their biomass (dry weight of organic matter in an area) in the form of extensive root systems or lignotubers or other large underground organs. Only about half of the living vegetation comprises aboveground shoots, and only a small part of that is in the form of leaves or other green parts such as stems in which photosynthesis takes place. It is therefore easy to see why production of new plant material is slow.

Some of the most complete information on scrubland productivity has been assembled from data compiled on Californian chaparral. While values derived from studies of scrublands in climatically comparable parts of France and Greece are similar, greater variation occurs in other shrub-dominated vegetation in less similar environments.

Chaparral 1.5 metres (4.9 feet) tall had a biomass of 46.2 metric tons per hectare, of which 16.7 was woody shoots, 3.7 leaves, 12.2 roots, and 13.6 litter on the ground. The annual accumulation of aboveground biomass in that system was 4.12 metric tons per hectare, of which 2.82 metric tons consisted of litter, with 2.64 metric tons of litter decomposing in the same period.

While such values typically demonstrate steady accumulation of biomass—in this case an annual aboveground increment of 0.13 metric tons per hectare—they do not take into account the occasional removal of aboveground parts through episodic disturbance, most commonly through fire. Productivity is more rapid during early stages of post-fire regrowth. For example, in one stand of French garigue, as the scrubland increased in height from 0.2 metres to 1.5 metres over a period from 1 to 10 years after a fire, its productivity declined from 4.7 to 3.2 metric tons per hectare per year.

Jeremy M.B. Smith