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Tropical Forests Introductory Article Tropical Forests Introductory article Sean C Thomas, University of Toronto, Toronto, Ontario, Canada Article Contents Jennifer L Baltzer, University of Toronto, Toronto, Ontario, Canada . Geographic Delimitation and Varieties of Tropical Forests . Historical Importance of Tropical Forests in Biology Tropical forests, while occupying only one-tenth of the world’s land area, are . Species Diversity disproportionately important in terms of global biogeochemical cycles and as home to . Alternative Mechanisms for Maintaining Diversity more than half of the world’s species. Research on these most complex of ecological . Tropical Forests in Global Perspective systems has focused on mechanisms for the maintenance of high diversity, and forest . Conservation and Sustainable Management of responses to fragmentation, logging, and other human impacts. Tropical Forests Geographic Delimitation and Varieties experience an annual dry period. In tropical dry forests of Tropical Forests (also called ‘monsoon forests’) the dry period is severe, and during this most trees drop their leaves in order to reduce Distribution patterns water loss. In semi-evergreen rain forests the seasonal drought is less extreme, and a leafless period does not occur To understand where and why tropical forests exist, one to the same extent. must begin with some essential facts of geography and Within these broad moisture regimes, tropical forests are climatic systems. By definition, ‘the tropics’ lie between subdivided on the basis of elevation and soil type, and 238N and 238Slatitude, the area within which the sun lies corresponding differences in forest structure. The distin- directly overhead at some point in its seasonal progression. guishing structural characteristics include canopy height, The flux of solar energy within this region is high, due to the crown layering and the presence (or absence) of different fact that incoming sunlight is projected at a 908 angle to the climbers and epiphytes. Tree buttressing, crown shape, leaf surface of the earth. This high solar energy flux results in a structure, and position of flower/fruit formation are other high rate of water evaporation over the tropical oceans and important physiognomic descriptors of tropical forests. of evapotranspiration over tropical land surfaces. The On a global basis the most important types of tropical result is a rising column of warm, moist air at tropical moist forest include lowland evergreen rain forests, upper latitudes. As this air rises it cools as a result of the gradient and lower montane rain forests, heath forests, peat swamp in air pressure through the atmosphere (adiabatic cooling). forests, freshwater swamp forests and mangroves (Table 1). Water condenses out from this air mass, generating rainfall. The air mass ultimately dries out, and is carried poleward from the tropics as a part of wind circulation Lowland evergreen rain forests patterns known as Hadley cells. At subtropical latitudes The tall, lush evergreen forests envisioned by most when near 23–308N and S, the now-dry air mass descends, referring to tropical forests are lowland evergreen rain creating a region of high pressure that corresponds closely forests. These forests are characterized by canopies with to the world distribution of deserts. As a consequence of multiple layers of vegetation and the presence of large these climatic circulation patterns, the earth’s equatorial canopy emergent trees. Lowland evergreen rain forests zone is warm and wet, corresponding to the broad band of generally have very high species diversity, with over 1000 tropical forests found along the earth’s equatorial axis tree species per square kilometre found in the richest forests (Figure 1). of Amazonia and southeast Asia. Canopy and emergent trees in lowland evergreen forests often have large Tropical forest formations spreading crowns with a radius of 420 m at maturity, can grow to more than 1 m in girth, and commonly possess The main types of tropical forest are distinguished by plank-like buttresses important in physical support. differences in the distribution of rainfall throughout the Beneath the upper canopy layer are smaller understorey year, by elevation, and by soil type. Tropical forests that trees, treelets, and a layer of herbaceous ground vegetation. experience ever-wet conditions with no month receiving Cauliflory and ramiflory are especially common among less than 100 mm of precipitation are generally referred to understorey trees in lowland tropical rain forests. One also as ‘tropical rain forests’, although a distinction is also generally finds abundant lianas; woody climbers that sometimes made between tropical ‘moist forests’ and ‘rain germinate in the understorey, but possess climbing forests’ in a strict sense that receive annual rainfall in excess mechanisms (such as tendrils or hooks) that allow them of 4000 mm. The two other main tropical forest types, to use free-standing trees as support structures. Lianas that ‘tropical dry forests’ and ‘semi-evergreen rain forests’, reach the canopy thus remain anchored to the forest floor. ENCYCLOPEDIA OF LIFE SCIENCES / & 2002 Macmillan Publishers Ltd, Nature Publishing Group / www.els.net 1 Tropical Forests Table 1. Key physiognomic features of tropical forest formations (adapted from Richards, 1996; Whitmore, 1998). Forest type Canopy Emergent Typical Tree buttresses Lianas Vascular Non-vascular height trees leaf size epiphytes epiphytes Lowland evergreen 25–45 m Common 45–180 cm2 Common Common Common Occasional rain forest Semi-evergreen rain 20–30 m Common 45–180 cm2 Common Abundant Occasional to Occasional forest common Dry deciduous forest 3–25 m* Absent 2–180 cm2 Occasional Common to Absent to Absent to abundant occasional occasional Lower montane forest 15–33 m Occasional 45–180 cm2 Occasional Rare Common Occasional to common Upper montane forest 3–18 m Absent 2–20 cm2 Absent Absent Common Abundant Heath forest 3–30 m* Absent 20–45 cm2 Occasional Rare Common Occasional Mangrove (mangal) 3–30 m* Absent 45–180 cm2 Prop roots and Rare Occasional Occasional pneumatophores common Freshwater swamp 3–35 m* Absent - 2–180 cm2 Prop roots Common to Occasional to Common Forest comon common abundant abundant Peat swamp forest 12–55 m* Absent - 2–180 cm2 Prop roots Rare to Rare to Occasional to common common abundant abundant common * Indicates forest formations generally characterized by gradients in canopy height and other physiognomic features. Also common are epiphytes; plants that live on other common. In many regions tropical dry deciduous forests plants (most often trees), but which at no point in their life grade into savannah ecosystems. history are rooted in the ground. Orchids, ferns and bromeliads provide many examples of tropical epiphytes, Montane rain forests which enhance tropical diversity immensely (epiphytes are An observant mountain-climber in the tropics will notice thought to comprise 10% of all vascular plants). pronounced changes in vegetation with altitude. The upper Another group of plants characteristic of tropical forests montane forests nearer the summit are short in stature are hemi-epiphytes, which germinate in the canopy, as do (1.5–1.8 m), with a canopy dominated by small, thick epiphytes, but produce roots that grow down the trunk of leaves affixed to gnarled branches. Bryophytes and the host tree to become rooted in the ground. The most epiphytes are extremely abundant in these forests, but important group of tropical hemi-epiphytes are figs (many buttresses, woody climbers and vines are virtually absent. species of Ficus), the fruits of which are an important The boundary between upper and lower montane forests is resource for many vertebrate species. generally quite distinct, with the latter showing a structure much more similar to lowland evergreen rain forest. Lower Semi-evergreen rain forests montane forests contain many features similar to lowland Semi-evergreen rain forests replace lowland evergreen rain forests. Changes in vegetation structure with altitude are forests in areas where there is an annual period of moderate ultimately driven by adiabatic cooling, though wind drought. As the name implies, some trees in these forests exposure and changes in soil characteristics can also be exhibit a seasonal leafless period, with the deciduous important factors. component comprising up to one-third of the upper canopy. Clumping of deciduous and evergreen trees does Heath forests occur, so some parts of the forest may be dominated Heath forests (called kerangas in parts of Asia) have a low, entirely by deciduous trees. Species diversity in semi- uniform canopy with little vertical stratification. The evergreen tropical forests is generally somewhat lower than relatively small leaves that are predominant in the canopy that of the evergreen rain forests. tend to be brownish and reddish in colour. The canopy is relatively open, and light levels in the understorey are high. Tropical dry deciduous forests An unusual feature of the heath forests are numerous In tropical dry deciduous forests (also called monsoon insectivorous plants, such as the Bornean pitcher plants forests) most canopy trees lose their leaves during the dry (Nepenthes) found in Asian kerangas. Podzolic soils season. The period of drought when trees are leafless is 2–6 develop in heath forests, which are highly acidic with very months. High light levels penetrate the understorey of good drainage. Water draining from heath forests is these forests for much of the year, and grass species
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