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ECOLOGICAL BIOGEOGRAPHY
Fong Lee Mo Kwan

WHAT IS BIOGEOGRAPHY? As the name suggests, biogeography is the geography of living things. However, it is more than just a survey of the geographic distribution of plants and animals; its subject matter covers no less than the vast reservoir of knowledge embodying the study of living things and their environments. Viewed in this way, biogeography is not very much different from ecology, to which it is very closely akin and from which many basic concepts are derived. But what is ecology? Ecology is a branch of biology environmental biology. The term itself takes a Greek root, for "oikos" in Greek means "house," and in the present context, it means ''environment." E. P. Odum has briefly reviewed that of the "biological spectrum" (protoplasm, cells, tissues, organs, organ systems, organisms, populations, communities, ecosystems, and the biosphere) ecology is concerned with the levels from populations and upwards.1 Thus put in the simplest language, ecology is the study of groups of organisms in relation to their environments. From the above description, it seems as though biogeography is no different from ecology at all. But, whereas the latter is mainly interested in the structure, the functioning, and the interactions of groups of organism within the ecosystem, the former is definitely more than that. As a part of geography, it has the same common interests as geography in that it is very much concerned with the place of man in the environment. Furthermore it is concerned with "all forms of relationship affecting the distribution, location and space organization of living things as they appear on the 'surface of the earth."2 Certain geographers,3 while recognizing that biogeography deals with the study of living things, plants, and animals inclusive, feel that it is
1 2 3 E. P. Odum, Ecology (Holt, Rinehart and Winston, 1963), p. 3. W. B. Morgan and R. P. Moss, "Geography and Ecology: the Concept of the Community and its Relationship to Environment," Annals, Association of American Geographers, Vol. 55 (1965), p. 340. K. C. Edwards and S. W. Wooldridge are the persons in mind and this idea is shown in the following works: K. C. Edwards, "The Importance of Biogeography," Geography, Vol. 40 (1964), pp. 85-97, S. W. Wooldridge and W. G. East, "Physical Geography and Biogeography," The Spirit and Purpose of Geography (Hutchinson, 1967), pp. 50-56.

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"more concerned with the world of plants than it is of animals."4 This is partly because animals, being more mobile, are less bound to their static habitats than plants are, and mostly because our plant world provides the basic supply of food for all animals, whether directly or indirectly, and so the study of plants forms the basis of the whole food chain upon which all other forms of relation depend. I would say that biogeography should be as much concerned with animals as with plants. The fact that research workers in this field tend to focus more of their work on plant geography is incidental, as at our present stage of advancement, we have a richer pool of information and data to draw from plant than from animal ecology. BASIC CONCEPTS BORROWED FROM ECOLOGY As mentioned earlier, many of the basic concepts used in biogeography are introduced from ecology. This in itself does not devalue the status of the subject, as cross-disciplinary communication and cultivation of knowledge are sound, healthy and scientific approaches,5 Thus a concise explanation of the basic concepts and terminology borrowed is indispensable, and it may even be appropriate to go back to some of the terms introduced in the earlier paragraphs here. Ecologically the term "population" refers to a group of oganisms of the same kind, whether they be ants, elephants or willows. Different groups of organisms (populations) living in a place are referred to as a "community," and the biotic community together with the abiotic environment make up the "ecosystem," as fish ponds, forests, and islands are all examples. The surface of the earth, or that thin contact zone of land, water, and air, upon which life prevails and different ecosystems co-exist is the "biosphere." Although the subject matter of ecology deals with the whole ecological spectrum, its basic unit of study is the ecosystem. The ecosysem is composed of four major constituents. 1) The abiotic substances are the basic matter, elements and compounds, that make up the environment, and air, water, and soil are examples. They form the raw materials of the habitat upon which the other constituents depend and from which the life cycles the different trophic levels evolve.
4 5 K. C. Edwards, op. cit., footnote 3, p. 85. E. A. Ackerman, "Where is a Research Frontier?" Annals, Association of American Geographers, Vol. 53 (1963), p. 433.

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2)

The producers are the organisms that can fuse the light energy provided by the sun with the abiotic substances of the environment to produce food, that is, organic matter. This magic power of transmutation the processing of simple inorganic into complex organic matters is possessed by green plants like trees, shrubs, and herbs.

3) The consumers are the organisms that depend on the producers directly or indirectly for food to build up more complex matter of their own, and they in turn may form food for higher consumers. These are the insects, birds, and animals. 4) The decomposers are mostly micro-organisms that break do\vn the complex protoplasm of producers and consumers alike and transform them back to basic matter, that is, abiotic substances. They are the fungi and the bacteria.

The green plants are referred to as the autotrophic components, or components that are self-nourishing, but both the consumers and decomposers, not being self-nourishing or autotrophic, are called heterotrophic ("othernourishing") components. The four constituents put forward may be structured into a hierarchy of two to five trophic levels (Fig. 1, Table 1). Thus in a woodland ecosystem, a sheep (herbivore) that feeds on grass may be devoured by a wolf (carnivore) which in turn may be consumed by a hawk (secondary carnivore), and if the hawk falls prey to bacterial or fungal decomposition, the chain breaks down, but the life cycle goes on and on, even though not along exactly the same line, it will be upon more or less the same pattern.
Table 1.The Structure of an Ecosystem

Constituents

Trophic Levels 1 Green plants 2 Herbivores 3 Carnivores I 4 Carnivores II 5 Microorganisms

Examples Soil, air, water grass sheep wolf hawk bacteria, fungi

I. Abiotic Substances II. Producers Autotrophic III. Consumers IV. Decomposers Heterotrophic Heterotrophic

Source: Compiled by author.

SHRIMP

MARINE ECOSYSTEM

TERRESTIAL ECOSYSTEM

FIG.I THE STRUCTURE OF A MARINE & A TERRESTIAL ECOSYSTEM

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The structure as noted above is classified, not according to species, rather, it is devised on a functional basis. The functioning of the ecosystem depends essentially on the flow of energy through it and the circulation of materials within it. The number of organisms and the rate of growth within each trophic level is therefore geared to this energy and material supply, and they also bear a chain effect to the well-being of its immediate upper level. Solar energy accounts for 99.99% of all energy available to the earth and it is upon this energy that all life depends. According to the first law of thermodynamics, this energy, which comes to us as an inexhaustible source from the sun, is neither created nor destroyed, though it can be lost to us by reflection back to space, or absorption by the earth and air. This light energy can be transformed from one form to another. The second law of thermodynamics states that this transformation, however, is not a hundred percent effective, for in the process energy is being converted from a concentrated into a dispersed form. This is the essence of the study of energy flow in an ecosystem. Of all the living things on earth, only green plants can convert solar light energy through photosynthesis directly as food energy, and heat energy is released in the process. According to Odum,6 of the total incident sunlight on green plants, noly about fifty percent is consumed in growth, respiration and food storage in the process of photosynthesis. As a large portion of the vegetal matter such as fallen leaves, stalks and roots are not used by animals, so the actual effective food energy available for the next level is only about one to five percent. This is what is implied in the inefficiency of transformation mentioned earlier in the second law. The channel through which food is transformed from one form to another (e.g. grass sheep wolf hawk) makes up the food chain. It becomes obvious from our description above that climbing up the trophic ladder, energy flow diminishes, less and less food is available, and fewer and fewer organisms can be supported. This decrease in number with ascension in trophic level is referred to as the biotic pyramid. It therefore follows that the amount of life (biomass) on earth is limited firstly by the amount of light energy available, and secondly by the effectiveness of energy transfer at each trophic level. Also, the biotic pyramid must generally range between two to five levels only, for beyond five, the actual
6 E. P. Odum, op. tit., footnote 1, p. 39.

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energy available becomes so small that life processes will be less effective. Odum made an attempt to quantify the energy flow in an ecosystem and the figures he quoted may well be used to clarify the explanations above (Table 2).
Table 2.Energy Flow in an Ecosystem

Incident Effective sunlight on photogreen plants synthesis

Effective food energy available for: Consumer I Consumer II Consumer III Decomposer Herbicore Carnivore 1 Carnivore 2 Microorganisms Sheep Wolf Hawk Bacteria 1% 10% 10% ?
15 1.5 .15

100% 50% 3,000 Kcal/m 2 / day 1,500

% each is a percentage of the figure preceding it. ? figures not available.

Kcal Kilogram calorie this is the amount of heat needed to raise 1 litre of water 1C Source: Figures based on E. P. Odum, op. cit., footnote 1, p. 38.

The circulation of materials from the environment through the organisms back and forth constitutes another function of the ecosystem. This movement is known as the biogeochemical cycle, and "bio" denotes life or biotic organisms, "geo" refers to the land, air, and water that make up the surface of the earth, and "chemistry" is the "science of the elements and their laws of combination and behaviour under various conditions."7 The elements present on the surface of the earth are numerous, but only about twenty are essential for the building of protoplasm. Some of these, like carbon, hydrogen, oxygen, nitrogen, potassium, calcium, magnesium, sulfur, and phosphorus and their compounds are needed in large quantities, while certain others are required in very small amounts. L. C. Cole estimated that of all the elements and their compounds required for bodybuilding, oxygen, carbon, hydrogen, nitrogen, and phosphorus account for
7 From The Concise Oxford Dictionary of Current English. (1952)

FROM VEGETATION

A
o
Ci

t-l
O C5
t*J O

'*-* 03

-\ f
PRECIPITATION

i
INFILTRATION

A \ 1 1
EVAPORATION

TRANSPIRATION

-a at

FIG. 2 THE HYDROLOGIC CYCLE

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over ninety-five percent of the mass of all protoplasm.8 They come from the soil, air, and water, and the greater part is taken up through the roots of plants as nutrients or via their leaves during respiration. These are passed on to animals as food for body-building. Some of these are returned to the soil through excretion, or to the air by respiration, but at death all the rest is returned to the environment through decomposition. The cycling of water on earth furnishes one such example (Fig. 2). Not only is the amount of materials present at each trophic level important to the efficient functioning of the system, but the rate of circulation through it, that is, the input and output of raw materials, is a deciding factor for its growth and prosperity. However, the proportion of the different elements required for body-building bears no relation to the proportion comprising the surface of the earth. Certain elements, like silicon, though present in large quantities on the surface of the earth, are required by living organisms only in trace amounts, others like carbon and nitrogen are required many hundred-folds percentage-wise than the proportion available (Table 3). Thus a more thorough study of the working of the biogeochemical cycle with an eye to conservation of the rarer elements for continuous circulation should be a first step in planning for land use and utilization of natural resources.
Table 3.Comparison of the Proportions of Different Comprising the Earth and Living Matter Elements

Elements Oxygen Carbon Hydrogen Nitrogen Phosphorus

Silicon Other elements

In the earth and atmosphere

46.00

In living matter
74.00 11.00 10.00 2.20

.03 .13 <.01 .13

28.00 25.00

.20 .05 2.60

Source: 8

Comparative weight (per cent) Figures based on L. C. Cole, op. cit., footnote 8, p. 88.

L. C. Cole, "The Ecosphere," Scientific America, Vol. 198 (1958), p. 88.

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Besides energy flow and materials circulation, the proper functioning of the ecosystem also depends on the conditions presented by the environment as a whole. To take an example, though sunlight is a must in any ecosystem, both too much or too little of it may be detrimental to life. In the case of a shallow pond with stagnant water and the tropical sun shining on it for twelve hours of the day, the abundant supply of light and the rise in temperature of the water promote growth, but the increase in chlorophyll and plankton leads to cloudiness and the decrease in oxygen in the water may be fatal to the fish in it. On the other hand, the absence of sunlight in the long winter season in an arctic pool will limit growth in both plant and animal life. The growth and perpetuation of life in an ecosystem actually depends on a complex interaction of different conditions. Any condition or any set of conditions that fluctuates beyond the tolerance limits, upper or lower, of any population so that growth either slows down or stops altogether in a system is called a limiting factor. Limiting factors may be climatic (such as temperature being either too hot or too cold, or humidity being too high or too low), or physiographic (such as the slope of the land being too steep or too gentle, or the elevation being too high or too low), or edaphic (such as the minerals, water, or air in the soil being too much or too little), or biotic (such as overwhelming competition between different species). These same factors, however, operating within the limits of tolerance may promote growth and productivity, and in this way, they are referred to as regulating factors. Thus the decline or well-being of a group or population in an ecosystem may be a biological indicator to changes in environmental conditions. Although the flow of energy, the cycling of materials and the physical environment all affect the biotic community either favourably or unfavourably, yet this is not a one-way influence, for the biotic components in an ecosystem, besides interacting upon one another, also react on the environment to bring about changes in time. Because of this reciprocity of relationship, we need to dwell more on this biotic factor. Every organism or group of organisms of the same species occupies a definite place and has a definite role to play in an ecosystem, and this place and role are referred to ecologically as the niche. Niche, however, is not habitat alone, it is in fact a functional classification of individuals in a community. Thus although the sheep of England, the bison of North America, the giraffe of Africa and the kangaroo of Australia occupy different habitats, they take the same niche in grassland ecosystems in different parts of the world, for they are all herbivores. In talking of niche,

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then, we actually look into the pattern of energy flow and the place in the food chain of a component. In a simple ecosystem, the energy flow is direct and the food chain is short, the removal of one niche in the system may restrict or even cut short the energy now and upset the smooth functioning of the system. Whereas in a complex system, the multitude of species in a single trophic level and the diversity of niches in the system build up a complicated food web, and energy flow is directed through different channels, so that even "with the reduction of any one niche, the metabolism of the system may well be absorbed by the other channels. A biotic community is very rarely completely static, it is continuously changing through time. As mentioned before, the flow of energy, the circulation of materials and the impact of the environment all help to bring about changes in the life of a community, but the biological processes of growth, development and reproduction in turn modify the environment and prepare it for future life to come. Thus an existing community is likely to be different from what it was fifty years ago or fifty years hence. Clements termed this process of change, or orderly, predictable community change from lower to higher life forms, succession. It involves gradual modification of the substrate and the micro-climate of an area through time so that new sets of organisms are better able to adapt themselves to the environment. In time this will lead to a climax community which is able to make the most efficient use of the energy and materials available and is therefore self-perpetuating, maintaining itself without much further change. This homeostasis or steady state represents a state of equilibrium between chemical, physical and biological processes. All the successional stages leading on to the climax are called serai stages. The sequence of succession from a pioneer community upon a newly formed surface where conditions of existence are unfavourable, as on a newly formed lava, onto a climax community is called primary succession. This is a very slow process involving over 1,000 years. When the sequence of succession of a community is interrupted, or when the original vegetation is destroyed, the new community that evolves establishes itself over more favourable ground for development than in the case of a pioneer community, and so the span of time leading to climax development is generally much shorter. Odum estimated that the return of a mature forest from a piece of abandoned cropland in a moist temperate area is in the region of 200 years.9
9 E. P. Odum, op. tit., footnote 1, p. 87.

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THE PLACE OF MAN IN THE ECOSYSTEM

Like the bees that drone in the fields and the quadripeds that roam the land, man too is one of the many components existing in the biosphere. The fact that he is a herbivore and carnivore all in one only gives him a double standing in the food chain, but does not change his nature as a component. In the early stages of man's history on earth, the simplicity of his skills and the inadequacy of his meagre tools made him subject to a greater extent to the controls of the environment. With the advancement of knowledge and mastery of increasing techniques and better tools, he is able to tamper with his environment more and more. He then becomes what C. O. Sauer called an ecologic dominant,10 or in other words, more a biotic factor than a component. In his attempts to change and dominate nature for his own survival, he has wrecked many ecosystems. Such tamperings are often not to his good, though not always to his immediate awareness; for instance, practising intensive agriculture may lead to soil exhaustion, or uncontrolled and indiscriminate burning may result in deterioration of grasses. The case of the westward march of the early settlers in North America is a typical example. The early settlers entered a rich continent virtually unexploited by man. In their eagerness to move on and to make fortunes, they cut down forests, brought the land carelessly under the plough and hunted ruthlessly for game. In a little more than two centuries, they had crossed over thousands of miles of virgin land leaving along their tracks large areas of gaping gullies, bare surfaces, and dust bowls. In these early days of ignorance their assumption of boundlessness was built on "a false philosophy of plenty and a myth of inexhaustibility."11 They speeded up what nature would have taken century upon century to create. This is only one case of man existing as a biotic factor destructive to the functioning of ecosystems. Recently man has gradually come to realize that in his attempts to understand the things around him his world and his biosphere it has escaped his attention to understand his own role in the biosphere. Man should not just extract himself from his ecosystem as though he existed apart as a director, controller, or observer, rather, he should thrust himself back, view himself as a part of the integrated whole and learn to coexist with the different components of the system. In other words, he should learn
10 11 C. O. Sauer, "The Agency of Man on Earth," Man's Role in Changing the Face of the Earth (The University of Chicago Press, 1965), p. 49. H. H. Bennett, Soil Conservation (McGraw-Hill, 1939), p. 33.

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to live and cope with nature in nature's way. This change of outlook is born out of necessity, that is, out of the awareness of the gradual depletion of resources and the destruction of nature. From this eagerness to coexist arises the need for planning and conservation, and the various attempts to bring about harmony between man and land, or the harmony of the functioning of the ecosystem. This harmony, which should be the first and most important lesson to learn in biogeography, is also the essence of the subject. DIFFERENT ASPECTS OF STUDY IN BIOGEOGRAPHY Regional Studies The basic unit of study in biogeography is the ecosystem. This has an areal expression, but its magnitude is elastic, for it ranges from anything from a small creek, a mound, an island, to an extensive forest. The essence of study is in the ecological energetics and functioning which can be measured and quantified. Studies can be made of different types of ecosystems so that the efficiency of each type can be compared. It is found that the net primary production12 of above-ground plant parts of different system are as follows:13 Tropical rain forest Boreal forest Mid-latitude grassland Tundra Desert 10.0 20.0 g/m 2 /day 2.5 .5 2.0 1.0 < .5

Comparison can also be made of the same type of ecosystem distributed over regions far apart. It is through comparative study that insights into the intricate functioning of communities are gained, and theories and principles which help towards planning and development can be drawn. We will have occasion to refer to this last point later on. In studying different systems, workers are confronted with the teasing problems of delimiting boundaries. In nature, life exists in association, not in isolation. Between one system and the next, there exists a gradient
12 13 Net primary production is "the organic matter stored in plant tissues in excess of respiration during the period of measurement." Adopted from E. P. Odum, op. tit., footnote 1, p. 39. These figures are abstracted from D. R. Harris, "The Ecology of Agricultural Systems," Trends in Geography An Introductory Survey (Pergamon, 1969), p. 137.

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upon which the dominance of one community fades into the next. Upon this gradient or transition, some criteria must be fixed on which one can demonstrate that the territorial units alongside the boundary are distinct. Arriving at this criteria is difficult, and even more so is the laying of this boundary line on the ground. The traditional delineation of boundary in most geographical studies has been on a single index such as that of morphology, climate or vegetation. This is unreal, for as mentioned earlier, life on the biosphere exists in association, and it is very much involved with the physical environment as a totality. Any line drawn based only on one aspect of this environmental link is bound to be incomplete. Regions should therefore be delimited on a functional basis for this functioning imparts reciprocity of all relationship and reactions. Once such functional units are derived, not only can we investigate the composition and working of individual communities, but we can also approach our study in terms of areal expression and spatial organization which are the key to geographical distribution. It is upon this successful delimitation of functional regions that the future of regional biogeography lies.14 Agricultural and Land-use Studies The practice of agriculture is actually the attempt of man to direct the energy of a natural system towards desired channels for the production of biota for subsistent or economic ends. This man-modified ecosystem, which is generally referred to as an agricultural system, may be conceived of in any scale from a tiny plot of farmland, a ranch or plantation, to a farm region of any size. Within this areal unit, the types of crops grown or livestock raised, the tools and labour (manual, animal or motor) used, the farming methods (ploughing, weeding, harvesting) applied, and the input (fertilizers and irrigation water) and output (crop) of material are all interrelated and form inseparable parts of the integrated whole. The study of agricultural geography is concerned with the various techniques of production and considerations of promoting economic gain, but biogeography investigates into the functioning of the agricultural system as a whole and more specially into the work of man in changing this natural biotic system and the effects of such changes. In nature, most of our environments represent generalized natural communities which are endowed with a complexity of life of many different
14 The main theme here is borrowed from R. P. Moss and W. B. Morgan, "The Concept of the Community: Some Applications in Geographical Research," Transactions, Institute of British Geographers, Vol. 41 (1967), pp. 21-22.

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forms. These are typified by a high diversity index15 with many ecological niches existing at all trophic levels. This results in a greater ease of adjustment with changing environmental conditions and a higher net primary production as all available channels are bridged and resources are used to the full. The tropical rain forest, which is tangled with luxuriant undergrowths, climbing plants' and tiers of trees of many species, and teeming with insects, birds and animals of comparable variety, is an example of a highly generalized natural community. With specialized communities, life is less varied and the diversity index is low. A direct contrast to the former type, it is also less productive and less stable. The boreal forest, with large stretches of comparatively uniform stands, is an example of this latter type.16 In his attempts to be systematic and to bring law and order to all his undertakings, man has applied many principles of simplification to the natural environment in his agricultural endeavours. Wherever he goes, he removes the existing flora and fauna and replaces them with one or a few types that he desires to raise to maximum growth and profit. History is studded with examples of man transforming grasslands into wheat farms, forests into plantations, and fertile valley tracts into paddy fields. Such transformation is in fact changing the generalized systems of nature into highly specialized ones. In so doing, he is leaving many niches empty and cutting short energy flow and materials circulation. In so doing, he has instituted a decrease in diversity of species, a decrease in stability, and a decrease in net primary productivity. One reason for this drop in productivity in most of our agricultural practices17 is that the photosynthetic mechanism is only operative for a few months of the year, that is. operative only during the growing season of a particular crop, and for the rest of the year when the crop is harvested,
15 Diversity index = cumulative number of species

16 17

" individuals counted Where the index is high, it means that the number of species is great and the number of individuals within each species is small. Where the index is low, it means that there is only a small number of species and the number of individuals within each species is great. D. R. Harris, op. tit., footnote 13, pp. 136-137. An illustration of this is in the annual net primary productivity of the following: Agricultural ecosystem Wheat, world average 344 gm/m 2 /year, Natural ecosystem Tall grass prairie, Oklahoma and Nebraska 446 gm/m2/year. Abstracted from E. P. Odum, Fundamentals of Ecology (Saunders, 1961), p. 73.

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the land lies fallow and productivity drops to zero. Besides, in arranging our crops in rows, we are restricting the leafy surface into a special pattern, leaving the parts of the ground between the rows and along field boundaries partly or wholly uncovered. This is very different from natural vegetation which is generally close-grown and exists in tiers. In practice we should follow nature's way of providing an assemblage of plants of a variety of species, growing to different heights and maturing at different seasons so that as much available energy is trapped as possible at all times. Odum has found that there is a direct relation between density of plant cover and productivity, and the yield of crop plants increases until the leaf surface is about three times the ground surface under the crop.18 In maximizing our ground surface for production, we should also seek to go along with rather than alter the niche structure of the natural vegetation by .substituting preferred species for wild ones. Thus an assemblage of fodder grass or root crops between commercial plants and fruit or other cultivated trees would bear close resemblence to a generalized natural system both structurally, functionally and spatially. For this reason, mixed farming is a more productive system than monoculture which is no more than a very highly specialized system. By extracting a member from a system, all the nutrients, or the concentration of energy and materials for body-building of that member are removed and this means an economic loss to the system. The principle to bear in mind in agricultural cropping for profit is to harvest without bringing degeneration or exhaustion to the system. Successive cropping of cultivated plants without renewing the soil will lead to soil exhaustion, decrease in productivity and deterioration of the crop. Excessive cropping of livestock will result in the lowering of soil fertility, deterioration and depletion of the grass, and degeneration of the stock, as the case of sheep rearing in upland Britain shows only too well some of these effects.19 Both, however, will eventually end up in soil erosion. Overcropping a member may lead to the extinction of a particular niche and this will impede the functioning of the whole system. In the case of plants, the problem may be solved by the application of fertilizers so that the materials exported are supplimented and the biogeochemical cycle is brought back to a balance. With animals, the control of the number of livestock on the ground should receive foremost attention in order to prevent the soil from being excessively trampled, and the sod from being completely removed, and furthermore, to allow the land to recuperate.
18 19 Ibid., p. 424. I. G. Simmons, "Ecology and Land Use," Transactions, Institute of British Geographers, Vol. 38 (1966), p. 21.

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Within the last forty years or so, farmers have been applying poisons of one form of another (D.D.T., diedrin, parathion, to name but a few) to their fields with the hope of eradicating pests. Some of the insecticides proved to be of immediate success, but the tragic thing is that they are also found to be effective is killing all insects irrespective of kind, the good and the bad, and those destined as well as those that are not. Another defect of insecticides is that, after being in use for a while the insects build up resistance and the poison has to be gradually increased in order to take effect. In the mean time, a lot of harm is being done to the life around, and the cumulative result is such that it brings pollution to the whole environment to the extent that it is even harmful to human existence. Rachel Carson20 attributed the insect problem to the intensification of agriculture. In nature, the varied forms of life coexisting in a community function as checks and balances to each other, so that no particular group or species is able to multiply and breed beyond the limits set by the biotic and abiotic environments. Yet with monoculture, or with clearance of immense acreage for the cultivation of a single crop or just a limited number of crops, these chosen species are able to thrive in abundance, and so also can the pests that feed on them. Here is an ironic incident of insect control21 which serves to illustrate that in his ignorance, man sometimes introduces harm to the environment which he dearly seeks to protect. In their attempt to destroy the cottony cushion scale which threatened the citrus industry in California, farmers there imported a parasite of the scale insect called the vedalia from Australia in the 1880's at great expense. Then in the 1940's, these fruit growers, like the other farmers elsewhere, tried the fancy, new, toxic chemicals on the fruit trees, and the result was that the vedalia were wiped out, and back came the scale insects. Conservation Studies From the realization that much of the resources of the world are exhaustible and that world population is growing at a rate faster than food production can catch up, and out of the desire to survive and to enjoy for the present as well as in the future the gifts of nature, there has emerged in recent years the recognition of the need for conservation. The term conservation has a no fixed definition and it bears a different meaning to different people. Used in the present context, it is the planned management of a community or a kind of resource so as to maintain continuous
20 21 R. Carson, Silent Spring (Houghton Mifflin, 1962), p. 10. Ibid., pp. 256-257.

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production without exhaustion or deterioration for the enjoyment and consumption of mankind. As the components within any community never exist in isolation, so conservation study, whether it be of a species, or a community, should take an ecological approach. With our preoccupation for getting a living, we have modified our environment without much thought. With time and with the consequences piling up, we have gradually come to realize that such modifications are not always best for community development from the ecological point of view, for reckless cultivation leads to soil erosion, uncontrolled lumbering leads to depletion of timber, lack of planning in watershed management leads to stream pollution and shortage of usable water, and many more such examples can be quoted. In order that we may evaluate objectively the nature and the direction of the changes brought about by such deformation, we should have at our disposal pieces of natural regions not yet contaminated by man for study. From such natural ecosystems, we can get an insight into the rate and nature of succession. Besides, by comparing the efficiency in functioning between natural and modified systems, we can better judge to what extent our tampering has been harmful or beneficial. It is through collection of information of this nature that we can decide what best to do in future. The designation of samples of ecosystems representative of the spectrum of natural regions of the world should be a first and urgent move towards conservation. Systems of aboretums, reservations and national parks have been introduced in different parts of the world with such objective in mind, but it is a pity that since man has inhabited the earth for the last few thousand years, such natural ecosystems have become more and more scarce. In the case of Peru, in an attempt to locate a place for use as national park in 1960, it was found that only two areas within the entire Amzon headwaters region were available.22 Huge volumes have been written on various aspects of conservation in connection with agriculture and soil, livestock and forestry, water and fisheries, as well as recreational areas. For an article of this brevity, the writer does not intend to go into these fields in any detail, but would like to stress the importance of taking an integrated approach in conservation. Conservation should not be viewed as a separate project for a separate department. To take water for example, it is the concern of everybody: the farmer is interested in getting an adequate amount of water during the growing season, the industrialist wants a continuous supply at all times for processing and driving the engines, the town-planner is concerned with
22 R. F. Dasmann, Environmental Conservation (John Wiley, 1968), p. 286.

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providing good, clean, drinking water for all, and the holiday-maker keeps an eye on water for recreational purposes. Each use may be in conflict with most of the others, and any policy formulated to cater for the interest of a specific group is bound to be at odds with all the rest. Water conservation then should be an overall policy accommodating the interests of all groups. Thus we need soil experts to advise on proper soil management to ensure efficient circulation of soil water and prevention of soil erosion, hydrologists to investigate into problems of water supply and demand, engineers to plan for the construction of dams and diversion channels for flood control and storage of water, agriculturalists for the proper management of farm land and irrigation water, biologists for the care and protection of wild life and natural vegetation so as to promote seepage, and other experts to look into matters of water pollution. These are all experts in their own fields, but pooling their efforts together they may be able to deal with the problem of working out a compromise to the benefit of all. To most people, the use of the forest is simply in providing firewood for burning, timber for building construction, and pulp for the manufacturing industry. For a knowledgeable few, it would include forage for grazing, and habitats for wildfire of different forms. Few would come to think of the aesthetic side of the forest in providing peace and serenity from the hubbub of life and areas for refreshment and recreation. Even fewer would realize the importance of the crowns of leaves and the mats of roots in bringing about soil and water conservation. The old concept of mining for timber is dying in many parts of the world, and accompanying this is the dawn of forest management. It is now realized that forest resources should be made forever renewable so that production and cropping should be balanced to maintain this goal. The diversity of expert knowledge and approaches in forest management points to the necessity for overall planning in conservation. Good forest management dictates that a detailed and up-to-date inventory of the available resources should first be made so that cropping can be planned in such a way as to allow maximum production, regeneration of the younger trees and also a balance in the whole system. The experience of German foresters23 in clear-cutting and replanting of pure stands of greatest demand since the 1840's has revealed that after two or three generations, the heavy drain of a handful of minerals had so degenerated the soil that the biogeochemical cycle broke down and production fell.
23 Ibid., pp. 192-193.

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Accompanying this was also a rise in losses due to pests and diseases. The introduction of a balanced system of mixed growth of many species and selective cutting since 1918 has shown that not only is production increased and quality raised, but there is general improvement of the forest lands. The type of trees that we grow is not the only thing that matters. Fire, insects and diseases are still the greatest hazards to forestry. Fire hazards can be prevented to a certain extent with care and should be the concern of all forest users. The institution of fire breaks, fire roads, and the introduction of systems of look-outs and fire vigilance are only secondary preventive measures. Diseases inflict a great deal of damage, and the most effective control measures should be sound forestry practices such as salvage by logging and isolation and the introduction of diseaseresistant strains of trees. Insects cause more damage than fire and diseases put together. In European countries, the promotion of forest hygiene24 in new growths of cut timber is found to be a promising way of defeating this last problem. Small animals, birds, ants, spiders and soil bacteria are all essential parts of a forest. In stands of young timber, the provision of nesting boxes can help to draw the birds and bats back to the forest and the work of insect hunting can therefore be taken care of both day and night. In Germany the red forest ants are found to be a very effective insect predator. Ant farms have been established and children are encouraged to go into the forests to distribute ant colonies. Spiders too are an indispensable member. It is estimated that a single wheel-net spider in her life of eighteen months may destroy an average of 2,000 insects. As the young spiders spin their nets in spring above the new shoots, they form a canopy over the crown of the tree thus trapping all flying insects coming from outside. In Newfoundland, the introduction of the voracious shrews had successfully wiped out in four years the sawflies which did great damage to the evergreen trees. The institution of chemical campaigns is likely to bring about a kill-one-kill-all effect, and is bound to end up in failure in the long run. It is only by guiding the forces of nature to our own ends that we maybe able to strike a balance, and that is why Odum advocated that we should aim at "conservation of the ecosystem rather than conservation of this or that species."25
24 25 R. Carson, op. cit., footnote 22, pp. 293-296. E. P. Odum, op. cit., footnote 1, p. 35.

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THE STATUS OF BIOGEOGRAPHY Right at the very beginning it was stated that the subject matter of biogeography is the study of that part of the surface of the earth upon which life evolves. The emphasis is on the spatial distribution of the different forms of life on earth, their interactions upon one another and upon the environment and vice versa, and the changes and successions that arise from such interactions. Central to this is the place of man as a part of and also as a modifier of the environment. This whole body of information on the inter-relationship of plants, animals, man and environment which is based on the ecosystem concept is also the main concern of geographers,26 and biogeography therefore holds a legitimate place in the discipline of geography. As the tendrils of geography stretch in different directions, as into geomorphology, hydrology and climatology on the physical side, into population and urban studies on the human side, and into marketing and transportation studies on the economic side, each of these off-shoots overlaps in one way or another with other disciplines of study and becomes a field of specialization of its own. The result is that geography seems to have lost its own identity and becomes more and more disintegrated. Biogeography is not just one of these off-shoots, it takes a happy unifying position between the living and the non-living environments, especially between man and land, and therefore possesses a binding effect in integrating physical, human and economic geography. As such, biogeography should provide a starting point for all beginning students in geography. In this world of rising population, dwindling per capita habitable space, increasing food demand, decreasing natural resources, and rising standards of living, the need for careful and sound planning of food production, land use, resource management, and improvement of the quality of living has become even more urgent. Although biogeography does not claim to be a panacea for all ills, yet the review of the various aspects of study in the previous pages does show that it has a utilitarian value and should therefore exist as an indispensable part of geography. Recent trends of study in biogeography have definitely taken on an ecological approach. Thus the study centres on the dynamics, the
26 This idea is summarized in Ackerman's statement that "an understanding of the evolution of space content as it is influenced by physical, biotic, and cultural processes must be an objective of geographic research." E. R. Ackerman, "Geography as a Fundamental Research Discipline," Research Paper No. 53, (Department of Geography, The University of Chicago, 1967), p, 19.

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structuring, and the functioning of the ecosystem. The ecosystem is a type of general system.27 It is an open system capable of leading towards homeostasis or a steady state, and is therefore self-regulating and selfperpetuating. It can also be conceived of in any degree of complexity and it therefore lends itself particularly to research and study. The use of quantitative methods allows precision of expression and easier1 comparison. Thus both the approach and the techniques used are sound and scientific. Biogeography should also be an indispensable part of a program of general education. The average man should be given a basic understanding of the world he lives in, an elementary knowledge of the interactions of living things and their surroundings, and the functioning of nature. - This would help to cultivate in him an ecological conscience which is so hopelessly lacking in most people and which is the key to the understanding, appreciation, and enjoyment of the beauty of nature. The preservation and the improvement of the environment is not the job of specialists alone, it is the responsibility of all who dwell on the land. Note: Though the term "ecological biography" coined by Moss and Morgan28 is adopted as the title of this article, throughout the whole of this paper, the full term never once appeared. The deliberate dropping of this qualifying word "ecological" is meant to avoid verbosity and undue repetition. The significance of the ecological approach has been explicitly clarified, so that the sole mention of biogeography should carry this ecological implication.
27 28 D. R. Stoddart, "Geography and the Ecological Approach," Geography, Vol. 50 (1965), p. 245. R. P. Moss and W. B. Morgan, op. cit., footnote 16, p. 23.