CLIMATE AND BIOLOGICAL PRODUCTIVITY OF THE TROPICS

The warm, wet, and relatively aseasonal climate of the tropics is apparently more favourable for maintaining higher diversity than anywhere else in the world. But why is this the case? One simple idea is that the high solar energy inputs and productivity of tropical regions result in greater numbers of species that can be supported energetically.


However, it is not entirely clear why a small number, or even one species, could not monopolize most or all of the incoming solar energy. Another idea is that climate stability is the main factor promoting species diversification and coexistence. In the harsher temperate and polar regions, species must be able to tolerate drastic fluctuations in seasonal temperatures. Species occurring in habitats nearer the poles are therefore adapted to a wider range of local environments in order to survive the winter months.

As a consequence, one expects a narrower range of adaptation to environmental conditions and narrower latitudinal and altitudinal distributions in the tropics, a hypothesis sometimes referred to as ‘Rapoport’s rule’. The more limited ranges of species in the tropics may allow for greater ‘species packing’ compared to temperate or boreal regions.

Although this idea has received much research attention, recent analyses give only equivocal support, at best, for Rapoport’s rule. On the other hand, it is clear that the relatively aseasonal nature of tropical rain forests allows for the evolution of highly varied and complex species interactions. This complexity itself contributes to the overall species diversity found in the tropics. For example, "dependent" ecological forms, such as specialist herbivores or predators, only persist in the community if their host species is present.

SPECIES DIVERSITY | Tropical Rain Forests Are More Biologically Diverse

Tropical rain forests are more biologically diverse than any other biome, lying at the extreme of a latitudinal diversity gradient that extends from the poles to the tropics. High species diversity in tropical forests is perhaps most impressively illustrated by the results of surveys of insects obtained using canopy fogging with broad-spectrum insecticides. 

A landmark study by Terry Erwin sampled the canopy of a single Luehea seemannii tree in Panama, finding 163 species of beetles (Coleoptera) co-occurring in this one tree. Through a series of extrapolations (based on estimates of the total number of world tree species and the proportion of insect species comprised of beetles), Erwin estimated that there may be 30 million species of insects occurring in the tropical forests of the world. This figure implies that 495% of the earth’s species remain to be described. 

While more recent sampling efforts have tended to yield lower estimates of tropical insect diversity, there are conservatively between 5 and 10 million species. This quantity is 5-to 10-fold greater than all species described to date. Thus, while tropical forests occupy only 7% of the earth’s land surface they are thought to contain over half of all of the species on the planet. The idea that species diversity of tropical animals (which are mostly insects) is a simple function of the number of plant species also implies that any effort to explain tropical diversity in general must, first and foremost, address the problem of the origin and maintenance of plant diversity.

HISTORICAL IMPORTANCE OF TROPICAL FORESTS IN BIOLOGY

Tropical forests have played a central role in the conceptual development of biology from the time of the major biological expeditions that began at the close of the eighteenth century.

Alexander Von Humbolt initiated the study of plant ecology on his voyages through South America in the early nineteenth century. While climbing Mount Chimborazo in the Andes, Von Humbolt characterized the vegetation changes with climate as he ascended. These early observations on plant distributions provided the foundation of the field of biogeography. The most significant development in biological thinking inspired by tropical forests was the theory of evolution by natural selection. Charles Darwin and Alfred Russel Wallace independently derived this theory as a result of their scientific voyages in the tropics during the mid nineteenth century. Darwin’s inspiration was his exploration of various parts of South America as the naturalist aboard the Beagle beginning in 1831.

Wallace conducted expeditions in both South America (1848–1852) and the Malay Archipelago (1854–1862), where he characterized two sets of fauna distinct to the different parts of the archipelago, a division now known as Wallace’s Line. For both Darwin and Wallace, the high diversity of species in the tropics, and particularly patterns of diversification associated with island groups, allowed an appreciation of evolutionary relationships not apparent in the poorer faunas of the temperate zone. The high diversity of the tropics continues to be an important inspiration and testing ground for ideas in biology, particularly in the fields of ecology and evolutionary biology.

LOWLAND EVERGREEN RAIN FORESTS

The tall, lush evergreen forests envisioned by most when referring to tropical forests are lowland evergreen rain forests. These forests are characterized by canopies with multiple layers of vegetation and the presence of large canopy emergent trees. Lowland evergreen rain forests generally have very high species diversity, with over 1000 tree species per square kilometre found in the richest forests of Amazonia and southeast Asia. Canopy and emergent trees in lowland evergreen forests often have large spreading crowns with a radius of >20m at maturity, can grow to more than 1min girth, and commonly possess plank-like buttresses important in physical support.

 

Beneath the upper canopy layer are smaller understorey trees, treelets, and a layer of herbaceous ground vegetation. Cauliflory and ramiflory are especially common among understorey trees in lowland tropical rain forests. One also generally finds abundant lianas; woody climbers that germinate in the understorey, but possess climbing mechanisms (such as tendrils or hooks) that allow them to use free-standing trees as support structures. Lianas that reach the canopy thus remain anchored to the forest floor.

Also common are epiphytes; plants that live on other plants (most often trees), but which at no point in their life history are rooted in the ground. Orchids, ferns and bromeliads provide many examples of tropical epiphytes, which enhance tropical diversity immensely (epiphytes are thought to comprise 10% of all vascular plants). Another group of plants characteristic of tropical forests are hemi-epiphytes, which germinate in the canopy, as do epiphytes, but produce roots that grow down the trunk of the host tree to become rooted in the ground. The most important group of tropical hemi-epiphytes are figs (many species of Ficus), the fruits of which are an important resource for many vertebrate species.

TROPICAL FOREST FORMATIONS

The main types of tropical forest are distinguished by differences in the distribution of rainfall throughout the year, by elevation, and by soil type. Tropical forests that experience ever-wet conditions with no month receiving less than 100mm of precipitation are generally referred to as ‘tropical rain forests’, although a distinction is also sometimes made between tropical ‘moist forests’ and ‘rain forests’ in a strict sense that receive annual rainfall in excess of 4000 mm.

The two other main tropical forest types, ‘tropical dry forests’ and ‘semi-evergreen rain forests’, experience an annual dry period. In tropical dry forests (also called ‘monsoon forests’) the dry period is severe, and during this most trees drop their leaves in order to reduce water loss. In semi-evergreen rain forests the seasonal drought is less extreme, and a leafless period does not occur to the same extent.

Within these broad moisture regimes, tropical forests are subdivided on the basis of elevation and soil type, and corresponding differences in forest structure. The distinguishing structural characteristics include canopy height, crown layering and the presence (or absence) of different climbers and epiphytes. Tree buttressing, crown shape, leaf structure, and position of flower/fruit formation are other important physiognomic descriptors of tropical forests.

On a global basis the most important types of tropical moist forest include lowland evergreen rain forests, upper and lower montane rain forests, heath forests, peat swamp forests, freshwater swamp forests and mangroves.

DISTRIBUTION PATTERNS TROPICAL RAINFOREST

The Tropical forests exist with some essential facts of geography and climatic systems. By definition, ‘the tropics’ lie between 23o N and 23o S latitude, the area within which the sun lies directly overhead at some point in its seasonal progression.

The flux of solar energy within this region is high, due to the fact that incoming sunlight is projected at a 90o angle to the surface of the earth. This high solar energy flux results in a high rate of water evaporation over the tropical oceans and of evapotranspiration over tropical land surfaces.



The result is a rising column of warm, moist air at tropical latitudes. As this air rises it cools as a result of the gradient in air pressure through the atmosphere (adiabatic cooling). 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 patterns known as Hadley cells.

At subtropical latitudes near 23–30o N and S, the now-dry air mass descends, creating a region of high pressure that corresponds closely to the world distribution of deserts. As a consequence of these climatic circulation patterns, the earth’s equatorial zone is warm and wet, corresponding to the broad band of tropical forests found along the earth’s equatorial axis.