continued . . .
"Like the cheetah with its speedy legs, the bees with their complex societies, and the elk with its prominent antlers, humans had developed their own trademark, which was aimed at enhancing their own security in an uncertain world."
"Control of food production was the most astonishing of all human
for buffering environmental disturbance." (Potts ,1996).
Fellow scientists have greatly misunderstood and scorned this mechanism of opposing forces. Yin and yang are not a part of western culture or thought. Termed group selection, ecologists such as Mc Farland (1993) understand it as claiming that "natural selection will favour behaviour that reduces the fitness of the donor if it benefits the group or species as a whole." By modelling perpetuity and compatibility I will show that the situation described by Wynne Edwards does arise. Instead of being of detriment to the individual, group selection only occurs when it is of benefit to the individual and the group. By this definition, then, group selection must include a direct benefit to individual fitness. "Group selection" is such a discredited and misunderstood term that redefining is necessary for the process that is taking place.
The most obvious mechanism by which animals "on their own "initiative", make density-dependent responses to changing conditions" is territoriality (Wynne-Edwards, 1986). Another form, social hierarchies, may be found in gregarious animals that form concentrations of animals such as in herds. Gregarious species may hold group territories and so the details can be intricate. However, the principle and possibility of density-dependent self-regulation still apply. A feature common to these various mechanisms is that they increase the intensity of intraspecific interactions.
An example of reproductive regulation associated with territoriality is the reproductive adaptation of lionesses. A characteristic of a species is generally an adaptation conferring some advantage, especially where it is important to reproductive success. If not, other strategies would rapidly evolve and replace it through natural selection. Strickberger (1985), a geneticist, recognises this fact when he says, "Because of selection over long periods of time, most populations may be considered to have achieved phenotypes that are optimally adapted to their surroundings." . . . "many phenotypes will tend to cluster around some value at which fitness is highest."
A lioness comes into heat every month or so when she is not pregnant. She is on heat for two to four days during which time she copulates once every 15 minutes throughout the day and night! This behaviour must have been selected for a reason other than reproductive success as the birth rate is still low. Researchers estimate that there are 3,000 copulations for each offspring that reaches adulthood, an extremely inefficient condition! (Krebs & Davies, 1991). Of the cubs born, only 20% reach adulthood. If numerical success in future generations alone provided a selective advantage, this strategy of the lioness could easily have been put at a selective disadvantage through a slight improvement in copulatory efficiency of some competing female or male! Clearly, it seems more efficient to copulate one or a few times instead of 600 times per born offspring! The economics of such behaviour simply does not make any sense.
In intraspecific interactions, as we will see below, it is behaviour conferring habitat stability that holds a selective advantage. Behaviour such as that in the lioness regulates or reduces the reproductive potential of the individual lioness and male lion (hear the alarm bells ringing in the evolutionist's mind). One must reason that another process must have selected for this behaviour that has the potential of reducing the number of genes transmitted to the next generation. It runs counter to the whole idea of the survival of the fittest.
Fitness has many dimensions. Fitness is achieved through natural selection, so relates the organism's genotype and phenotype to the environment. An individual fit in one environment may be totally unfit in another. Fitness is relative in that we measure it as the contribution of genes to the next generation compared with other genotypes. An organism can promote its genes in the next generation through reproduction and assisting in the survival of relatives with the same alleles. They term the former individual selection and the latter kin selection (Beck et al, 1991). The aspect of fitness to emphasise is its relation to the habitat of the organism - its ecofitness.
There is a very fine line of interpretation that is now to be crossed. This concerns the idea of "survival of the fittest." Creatures such as cheetah exhibit territoriality, even killing intruders. On the one side of the balance is the belief that this represents survival of the fittest with the following reasoning: "the strongest individuals are despots, grabbing the best quality resources and forcing others into low quality areas or excluding them from the resource altogether" (Krebs & Davies, 1987). Through healthy competition, only the strongest survives and reproduces. Those able to hold territories are more able to pass their genes onto the next generation as:
 they can get mates,
 they occupy the better habitats with more food.
The cheetah is a good example here of an animal lowering its reproductive potential (but not its survival potential) through its behaviour. It has the potential of producing many more offspring than it does in any season. Its territorial behaviour limits the number of offspring it produces as an ecological necessity, although a greater number of offspring produced in any season is a genetic advantage. How does territoriality achieve this?
Some male cheetah hold territories for periods of four to six years, while others are wanderers. On the Serengeti plains, up to 40% of males are solitary wanderers without territories. Adult female cheetahs are usually solitary, but their cubs remain with them for at least one year. Sibling males often pair up in twos or threes as these unions provide greater success. Where cheetahs thrive, such groups are bigger and more common. More than one male may cooperate to establish and hold territories, with such coalitions being more successful. Territories are marked daily with sprayed urine or prominent trees and rocks. Cheetahs attack, maul and sometimes kill intruders. When a female comes on heat (oestrus), males detect this through the smell of her urine and territorial behaviour intensifies. Fighting among males increases and the males follow the female closely. Individuals slap each other with downward swipes of their front paws and may bite each other. Sometimes they kill a combatant. Courtship lasts two to three weeks before the female becomes receptive. By this time a dominant male mates with the female.
The drive to perpetuate a species numerically we can call simply and literally perpetuity. A cheetah occupies a particular niche and habitat. They have an ecological role and dependency. Their size, behaviour and physique make them specialists. On the Serengeti Plains, the Thompson's gazelle forms about 80% of their diet. Generally, their preferred prey is antelope weighing up to 60 kg, but they also predate upon ground-living birds, young giraffe, hares and porcupines (Skinner & Smithers, 1990). Many other animals are too big, such as adult giraffe, buffalo or eland, others are usually too elusive, or do not suit their hunting style, such as red buck, dik dik, birds and tree dassies. Creatures such as baboons and adult warthog are quite dangerous.
The question must be how territoriality is selected for, or more specifically, what forces have led to territoriality in cheetah? Most say it is this bloody nail, tooth and claw competitive battle for dominance by the strongest. "In one way or another the limitations on the density of a population ultimately work through the intraspecific competition that results when a number of animals utilize a common resource that is in short supply. Of the various components of the environment, the two that are apt to be in short supply in relation to population density are space and food. Competition for them may result in aggressiveness, stress, starvation, or emigration, but in the last analysis some element of this competition is involved" (Smith, 1974). This is the "resource limit interpretation".
Let me now place some weight on the other side of the balance -
ecofitness. The natural habitat provides the conditions for
natural selection to cause another form of life perpetuating behaviour:
interdependence and co-operation termed compatibility as opposed to
competition. In this context, territoriality is a response selected for
by the "need" (a necessary condition for survival) to maintain the
stability of the habitat or ecosystem upon which the animal depends.
Territoriality is a behavioural strategy that enhances habitat
stability and excludes the "cheaters" from an area big enough to
maintain the habitat. It reflects the animal's strategy
to regulate its population so that it does not detrimentally or
significantly alter the habitat
to which it is adapted and upon which its survival and the survival of its
offspring depends. Enter the holistic domain of ecofitness. This "need" is the
behavioural strategy that confers habitat stability and thus maximises
potential within the habitat to which the animal is adapted. This is
"habitat stability interpretation". When birds can evolved
complex migratory or nest building patterns and individuals of a single
ant colony can evolve separate physical forms, then organisms can also
evolve mechanisms that prevent habitat modification.
Behaviour is as much subject to selective forces as is form. Garter snakes (Thamnophis elegans) of the SW United States are largely terrestrial slug eaters on the coast and aquatic underwater feeders inland, catching frogs, fish and leeches. Wild-caught inland snakes do not eat slugs. Tests on new-born snakes showed that 73 percent of coastal snakes readily ate slugs, while only 35 percent of the inland snakes did so! Crossbreeding showed that the difference was heritable and therefore genetic, with the offspring having an intermediate preference for slugs (Krebs & Davies 1987).
An organism's survival depends as much upon the perpetuation of the
habitat to which it is adapted and the niche that it occupies, as it
depends on individual
fitness and reproductive success. Here we have yin and yang in nature,
conveniently called ecofitness! We
should define individual fitness in terms of the animal's niche. An
is only fit within the context of its habitat, as shown by the two
of garter snake. The animal depends on the unchanging consistency of
habitat, or at least a rate of change to which the animal can adapt and
evolve. Unrestrained numerical success as progeny into the next
generation on its own will destroy the habitat.
Many animals preserve habitat stability by the simple and common
strategy of territoriality. Convergent evolution has led to this
strategy repeatedly in a whole range of life forms and from the very
beginning of life. Cheetah males maintain territory sizes selected for
through each generation and
all their history as a behavioural adaptation to their habitat. This
territory encompasses its prey and is of a size sufficiently large to
its prey. Females may hunt within this territory and so be available
they come into oestrus. Chasing away conspecifics conserves this
but also puts these dislodged or displaced cheetah at a disadvantage.
Naturally the fitter individuals maintain territories and so has access
to female mates. Feedback from the habitat of the cheetah responds
sensitively to dynamic changes
in the cheetah population. If the behaviour of the cheetah increases
numbers in the next generation, it may deplete the resources upon which
it depends, starving itself and reducing its offspring's viability over
the long term. Immediate and subtle changes in the habitat of a
in response to the behaviour of the species, serves as a form of
and selective force, regulating species numbers. This feedback force is
If the cheetah depletes its food resources or alters its natural
within its territory, its own activities reduce its survival potential.
is a (passive, non-teleological) response by the habitat to the
of the cheetah. Survival depends not only upon numerical perpetuation
the species, as man sees it today, but also compatibility with the
Generalising about behaviour never provides the full picture, which is always more complex. Male cheetahs use urine and faeces as territory markers (Skinner & Smithers, 1990). However, these markers are only effective for 24 hours, so another group may move in and use the same area. This allows considerable overlap between home ranges. Males may form bachelor groups of up to five individuals. Female urine has no territorial significance, but they do occupy home ranges and are solitary. Females hold areas, but it is uncertain if they drive away other females. There is no doubt however that the behaviour of the males and females spaces individuals throughout their habitat more than if they were randomly spaced (the null hypothesis). Contact between groups usually leads to nothing more than threatening behaviour. Individuals interact with ears drawn back, the head held low and the mouth open. This aggressive behaviour results in there being fewer individuals per unit area than the habitat can support, so minimising their impact on the ecosystem. Cheetahs are keeping their numbers below the carrying capacity of the habitat through their own behaviour. As it is the fittest individuals that hold territories and produce the offspring, these individuals are effectively ensuring their perpetuation by limiting their own numbers through territoriality. This behaviour has a holistic effect, as it is in the interests of individual, group and habitat survival. It is the result of Darwinian natural selection.
Returning to the lioness discussed above the male sexual drive is selected for maximal perpetuity. If his mating success as mature offspring was more successful than one in 3000 it would threaten its habitat. Selection acts upon the female so that the rate at which she falls pregnant maximises the survival potential of the offspring produced - this requiring the optimal physical condition of the female and a stable habitat. The solution is a low pregnancy rate in relation to actual copulations, but one suited to their natural habitat, niche and social structure. This apparently inefficient mating behaviour makes sense within the context of the animal's habitat and physiology.
A similar condition is found with humans: "whatever the main biological function of human copulation, it is not conception, which is just an occasional by-product" (Diamond, 1992). Human females do not know when they are ovulating (and nor do the males). In many other primates, the area around the vagina and in some the buttocks and breasts swell up and turn red, pink or blue to advertise their condition, while the females become more receptive. Our closest living relative, the chimpanzee, shows very different mating behaviour to humans. A female is fertile and receptive for four to six weeks during which she mates on average six times per day. Her genital area becomes swollen and enlarged and is pink or red to advertise her condition. She mates with any male, but as ovulation approaches, the dominant males guard her and fight to mate with her. After ovulation her genital swelling subsides and she will not be sexually receptive again for three to four years (Palmer et al, 1994). The closely related bonobo has a different social structure with the females dominating the groups.
Human females are receptive with or without concealed ovulation, so with maximal sexual encounters there is only a 28% probability of conception (Diamond, 1992). This fact may give some clue to primitive man's social organisation and ecological role, yet why humans and chimpanzees have evolved such different sexual strategies is not clear. Continual receptivity by human females may have evolved to maintain pair bonds and male attendance that became necessary to survival in a harsher environment. Nomadic hunter-gatherers will need to carry young children when on the move. This places a limit on the frequency at which a female can fall pregnant. The spacing between pregnancies in nomadic women is about four years while sedentary farmers can have children at shorter frequencies of about two years (Diamond, 1992). Interestingly the parallels between lions and humans go further in that it was the most widespread mammal after humans 10,000 years ago, being found in Africa, much of Eurasia, North America and northern South America.
A habitat responds to the individuals interacting within the habitat. If the cheetah eats all the gazelles within its territory, it must move in search of more food and fight with other territory defending cheetah. Generally this will put this cheetah at a disadvantage. It cannot mate until (and if) it can establish a new territory. Such an individual will be maladapted to its habitat. These habitat responses maximise the survival potential of individuals of the species whose behaviour confers habitat stability through territoriality. Cheetahs ensure habitat stability through defending a territory, so preserving its food resource. Territoriality provides an intense interactive mechanism that animals utilise to sort out the fitter of the population that will mate and contribute to the next generation. This in turn controls the gazelle population, preventing overgrazing. Some antelopes (bovids) are territorial, so preserving their own habitat. Interdependence between species of a habitat evolves because of such interactions.
Considering ultimates, the less influence an animal has on a habitat, while still able to survive, the greater its survival potential and thus compatibility or ecofitness. This idealistic condition requires that even as the habitat evolves, the animal does not diminish its survival potential. As it depends on a food resource, this is not practical. However, for a species such a man with diverse dietary possibilities, a habitat could evolve without affecting his survival, as with the Bushmen hunter gatherers and perhaps early man. They used all habitats without modifying them and thus did not significantly affect ecosystem structure.
It is significant that the ancestors of man, bipedal apes appear to have arisen in the Great Rift Valley when dense rain forest gave way to a mosaic of environments, from forest to woodland to shrub to grassland. The formation of this valley through geological and tectonic processes produced many ecological barriers and micro-environments (Leakey & Lewin, 1992). In this dynamic environment a bipedal creature evolved and through necessity, had to be able to use a diverse range of habitats. As a hunter gatherer, not specialised or restricted to any specific habitat, early man had little impact upon his environment. Early man was thus adaptable and through coincidence compatible. This natural occupation of a pinnacle position in ecosystem processes, although through an accident of events, propelled this creature to the position of ecosystem dominance. This creature could move across and between ecosystems and survive, so was not constrained by or subject to any ecosystem. Humans remained dietary generalists. Today, intelligent man is being forced through sheer force of numbers and success as a species to recognise the necessity for the compatibility that was so naturally practised previously.
If we now try to define compatibility we arrive at a problem. We cannot numerically quantify it. We could allocate to it a relative figure from -1 through 0 to +1. Thus, at zero the species does not influence habitat or ecosystem structure. At +1, it is having maximal positive influence in some manner and at -1, destruction of the habitat is maximal. Primitive hunter gatherer tribes exist at a position close to 0. Today our influence as modern technological and civilised man is negative. The future objective should be to transform this trend into a positive process through the development of more benign technologies.
We can develop this concept in another direction. Lions enlarge their territory when food is scarce and when plentiful, reduce its size. The carrying capacity for the environment of such a predator depends upon the number of prey per unit area. Available grazing and the productivity of the habitat limits prey numbers. Many abiotic factors such as soil type, rainfall, temperature, sunshine etc., determine productivity. If a hypothetical lion and lioness - a pre-territory-era lion pair - act in response to the innate or instinctive drive to perpetuate to numerically maximise their genes in the next generation, what selective pressure, constraint or limit will they most rapidly encounter? If in each generation they produce six surviving young, the area will rapidly become overpopulated with lions. (First generation, one male, one female, 2nd generation three males and three females, third generation, nine males, nine females, fourth generation, 27 males and 27 females . . . ). The many offspring will eat all their prey and a population collapse will occur. Evident here is a very strong and rapid response from an ecosystem to the presence of a species. This acts as a selective pressure. What I am saying is that the biotic component of the ecosystem provides a direct selective pressure or response as negative feedback to the interacting species. An animal such as a lion continually exploiting its habitat to maximise its number of offspring may even cause a food shortage so that its offspring starve. Territoriality is one solution, found in many creatures, as a response to negative feedback from ecosystem constraints. In nature, ecosystems full of populations of interacting species have been evolving throughout all time. As the species have evolved, so the ecosystems have evolved, a principle of complex, holistic systems. Ecosystems respond to the presence of the inhabitants. This feedback acts as a biotic selective force or constraint to which the animal must adapt through behavioural, physiological or physical (morphological) changes!
What is the effect of territoriality according to this perspective? The territorial animal maintains an area big enough to feed itself and its young and thus perpetuate, without depleting its food resource - forming a core or unit of (biotic) stability. Behaviour is selected for which maintains the stability and perpetuity of the ecosystem and associated species. Abiotic environmental fluxes remain density independent and have to be adapted to as necessary. The maintenance of a territory assures that offspring over many generations will have food if cyclic abiotic influences remain stable and predictable. This "if" is the main cause of extinctions of species adapted to specific ecosystems to the extent that they are not able to cope with change. Non-territorial conspecifics are excluded by the territorial behaviour and so they simply eliminate the problem of "self-seekers" or "cheaters" in this context. Only the fittest maintain territories in each generation and so breed. This concept of territoriality conferring habitat stability is to be found in theoretical ecology texts (Begon, et al, 1986) and termed "self-limitation". Ecologists term this intraspecific competition or mutual interference.
In this book, such processes are described with the MELV formula. Predators that hunt "efficiently", so depleting their prey, attain a high equilibrium density, but are less stable, with more persistent oscillations in predator and prey numbers. As prey are depleted predator numbers stabilise or collapse. Too many prey may degrade the habitat, leading to a chronic decrease in prey and then predator numbers. An "inefficient" predator with many prey in relation to predators also shows oscillations, but these are damped and equilibrium prey density approaches the prey carrying capacity. With strongly "self-limited predators", stability is high, often without oscillations. Predator numbers would tend to be low and prey numbers will be near the carrying capacity, but both populations will be stable (Begon et al, 1986). Thus a compatible animal, which exhibits (self-limiting) behaviour that reduces its effect upon the habitat upon which it depends for survival, confers stability to that habitat. This is an established fact in standard ecological texts. What is significant here is the recognition of conferring of stability to the system and that the prey remains at the carrying capacity with predation having minimal effect.
The most important consequence of territoriality is population regulation. Often, displaced individuals without territories fail to reproduce or have a lower reproductive success in sub optimal habitats. Such a situation was found in great tits. Successful territory holders occupied woodlands, while displaced birds occupied hedgerows where reproductive success was lower. Hedgerow occupants would rapidly occupy a vacated woodland territory.
The traditional view of this situation is from the individual perspective, with contest, winners and losers (Begon et al., 1986). Wynne-Edwards (1962) promoted an alternative, that the regulatory consequences of territoriality must be the "root cause" or selective force resulting in the evolution of this behaviour. However traditionalists did not understand him or perhaps he did not describe the mechanism properly. They rejected his ideas in favour of the need to describe natural selection in terms of some advantage accruing to the individual (Begon, 1986).
Wynne-Edwards promoted the idea that the population as a whole benefited from the territorial behaviour as the population did not overexploit its resources. Other professionals defined this as group-selection and they rejected the idea for "fundamental reasons". Evolutionary theory could not provide a mechanism for group selection. Natural selection as defined according to traditional theory could not explain the mechanism behind apparent group selection. Traditional territory explanations speak in terms of energetic costs and benefits, little recognising the tautology in this explanation: just as surviving animals survive, so energy gained in holding a territory must exceed the energy cost required to defend the territory. If not, the animal would not exist. What the researchers discover is necessarily true if the animal is to live (Beck, 1991). Similar to the mutually exclusive God-or-evolution arguments, ecologists claimed that the net advantages accruing to individual competitors cannot also be those required by the group. A change of emphasis is necessary from individuals and groups to individuals and their habitat.
Further consideration of this interaction raises the possibility that a compatible predator not only confers stability through not overexploiting its habitat, but may have a positive and beneficial effect upon the habitat. If prey numbers are close to the prey carrying capacity for the habitat then the predator must be removing the least productive components of the prey population. Predation is selective, taking the old, young, sick, injured, vulnerable or weak as these are the most easily caught. This confers increased fitness to the prey population and allows more dynamic growth within the population, especially with a compatible (self-limited) predator. A second important factor already mentioned is that the predator prevents the population explosion of the prey in many cases and so prevents the destruction of the habitat vital to both prey and predator.
Compatibility is an adaptation to reduce the species effect
the ecosystem, so maintaining ecosystem stability. Creatures are driven
the impulse to perpetuate, but as demonstrated, this cannot function
In complex ecosystems it is found associated and inseparably linked to
the necessary condition of compatibility with the system, conferring
stability. Compatibility and ecofitness is the result of a selective
force due (negative
feedback) to the influence of the species interacting with the
It is the result of the coevolution of species that encounter a
of constraints in their environment. An animal destroying its habitat
lowering its survival potential or ecofitness as the animal depends
the ecosystem for its existence. The animal's struggle for survival is
the context of the biological system with which it, by necessity, has
coexist. A species has to be compatible with its ecosystem and we
many amazing natural relationships as an expression of this
the diversity of which represents the diversity of possible
that have evolved. These ecosystem constraints act as a force creating
natural compatibility found in nature! It is this incredibly simple
that provides the holistic mechanism and holistic nature of ecosystems
by so many people.
"Fitness" now takes on a new meaning - there is a component that can be called ecofitness. Ecofit animals are compatible with the natural habitat to which they are adapted, so at best it is beneficial as it confers stability to the ecosystem. Here, we have an almost mathematical-type formula. An animal can have a negative influence upon the stability of an ecosystem; it can have no effect upon the ecosystem at all; or it can affect the ecosystem in a positive, beneficial way. Modern technological man is having a gross negative effect upon global ecosystems, to his ultimate detriment. African Bushmen who lived in Africa for aeons before the arrival of the black Bantu African and white Europeans' colonisation of southern Africa appeared to have no effect upon ecosystem structure - being absolutely compatible. A future interactive solution to the current destructive chaos is for man to become central to ecosystem processes and positively influence ecosystem structure. An alternative is the doomsday scenario and our end.
With predator-prey interactions seeing the benefit of compatibility or self-regulation is easy. In interspecific interactions a community of associated organisms, the benefit of compatibility is less obvious. This has led to the development of anthropomorphic ideas such as competition. In the 1970s and 1980s ecologists believed that interspecific competition was important in forming community structure. They have now modified this to include "non-equilibrium and stochastic factors, such as physical disturbance and inconstancy in conditions" (Begon, 1986). Ecologists have formulated the competitive exclusion principle, which says that "if two or more species compete for the same limiting resources, then all but one of the species will be driven to extinction" (Begon, 1986). The law of compatibility takes the same observation and turns it from one of competition to one of benefit. If two or more species interact, utilising the same limited resources, then natural selection will favour the evolution of interspecific interactions with less mutual effect than intraspecific interactions. Organisms adapt to the conditions and constraints of their environment. Solutions in nature are necessarily economic as greater efficiency can be a selective advantage. This economic interaction can be expressed metaphorically, and mathematically in a simple model.
First metaphorically, consider that a community of people has an average interactive effect of one, suggesting that statistically, the average effect or intensity of interaction upon one another is equal. This value of one represents the (statistically) average amount of energy required for intraspecific interactive activities converted into a relative coefficient. Nearby is a herd of wild horses that graze free. Their intraspecific effect is one, while their interspecific effect from the human's is 1.1 as chasing the horses away from the plains, reserved for grazing by his domestic herds, costs the man more on average than human-human interactions. A man-horse interaction thus "costs" both species as is found in classic competition. However upon domesticating the horse, the man can travel greater distances and carry larger burdens. The horse in turn receives intelligent care, protection and a wider distribution, so has enhanced survival through the new interaction. For both, the interactive values (or effect or cost) becomes 0.9. In other words both the horse and man are more beneficial to each another than a man-man or a horse-horse interaction. Instead of competing for a resource that both needs, the plains for grazing, the association shifts to one of interdependence. The two animals thus become compatible and of benefit to each other! A coevolutionary process has begun. Their interactive intensity has changed from a costly energy expenditure to a beneficial interaction. We see this in nature as interdependence between species of an ecosystem.
Can anyone deny interdependence within ecosystems? This interdependence confers ecosystem stability. It evolves through natural selection because of the association of organisms. What I am saying is that when many species interact, perpetuation of the species does not result in the evolution of interspecific competition, but the tendency for the evolution of interspecific compatibility, so that interacting species have less effect upon each other than do the average of intraspecific interactions. Ultimately the energetic costs or intensity of intraspecific interactions must drive the process as fitness is a relative value. Biologists find many examples of this process. Two tree frog species, Hyla ewingi and H. verreauxi have similar calls where their ranges are separate, but where their ranges overlap, selection favours the evolution of greater differences between the calls (Gould & Gould, 1989). This process of reducing the interactive effect or intensity of interactions is selected for as it leads naturally to:
 Improved survival potential of each species as they waste no time in faulty communication;
 Ecosystem stability through population stability;
 Fewer extinctions than would be witnessed in aggressive interspecific competition.
 It is energetically more efficient and so selectively superior .
This process is not easily evident as we cannot witness evolution but only the result which is the interdependence witnessed in nature. We cannot measure the interactive costs of dinosaurs from 70 million years ago and compare them with modern animal life to show that modern life is more efficient energetically. A comparative study of vertebrate anatomy through time will reveal improved efficiency in form so substantiating this hypothesis. Through reverse engineering, we can trace reduced robustness and improved engineering design through time.
Models of the evolutionary process can mimic competitive, neutral and compatible interactions over time to see what the effect is. We must recognise the many constraints upon natural systems, such as intrinsic growth rates, environmental carrying capacities and predation rates. Biological systems are full of natural constraints and different associations. Humans for example cannot metabolise more than 5,000 calories per day, there is a limit to radiant energy reaching the earth each day, flowers use insects to effect pollination and so forth.
I illustrate this process through the application of the modified energetic Lotka-Volterra model (MELV model). One has to ask whether the results of the model illustrate intrinsic constraints of the mathematical model or display intrinsic, natural processes. The weight of evidence seems to suggest that the mathematical model reflects a real process in a general way so that the mathematical outcome illustrates the natural outcome. We will develop and investigate the MELV model.
Note that this approach is not anti-technology. Technology needs to play a central role in providing economy, efficiency, convenience, ease, dignity, care and so forth to our lives. We need high levels of efficiency to sustain high population densities. We must either rescue nature by the application of technology or return to simpler, subsistence lifestyles at a much lower population density. Our skill in managing technology will decide which route the forces of nature will allow. Nature is testing us at this very moment.
Thomas Hobbes, a writer from the 17th century described a good law as "needful, for the good of the people, and withal perspicuous." Well, today we need to go further and establish laws and principles needful, for the good of the earth and all its species and resulting in clearly beneficial natural responses. Man's interaction must no longer be irrational, violent, destructive and unpredictable. Anyone who observes nature can detect the harmony present. Science has revealed intricate relationships and interdependent needs between groups of organisms. Humanities' association with nature requires that he restore the necessary compatibility with nature. Success is not to be measured as numbers alone, as we are discovering, but ensured perpetuity requires compatibility and not competition with nature. We need to instil a new social consciousness. The gross influence of billions of people upon the earth is forcing us to realise the need for compatibility. Economists call this need sustainable development.
A new relationship needs to be found with humanity at the centre of ecosystems. We need to establish the formula for perpetuity and compatibility. We need to define humanity's role. This is going to require a new awareness and behaviour among individuals. If we multiply each of our actions by a million, we can see that collective changes can have a very large effect. Governmental priorities are going to have to change. Economies of governments are going to have to shift away from finite resources such as gold to an emphasis upon a sustainable natural productivity and compatible technology. Social structures will need to elevate each individual to a high level of education so that people appreciate intellectual art as is visual, sensual and musical art today. Industry needs to adapt urgently through the production of biodegradable and recycled products.
A full definition or understanding of compatibility and perpetuity
will incorporate the idea of sustainable development. Implementation of
sustainable development formed the central thesis of the 1992 United
Nations Conference on Environment and Development and yet at the end of
this conference, this idea still lacked complete definition and thus
understanding. Thus, right now, "researchers are seeking pieces of a
mosaic of development, the overall pattern of which, like the web of
biodiversity, can only be guessed at" (Halloway, 1993). Food and energy
production needs to be localised and diversified
to reduce expensive transport costs and minimise disease problems
associated with monoculture. Technology must advance far in the
direction of electronics and efficient communications as travel becomes
more expensive, unless the science of nuclear fission is mastered or
they discover another pollution-free source of energy. We must better
respect nature. Statisticians could play a leading role in predicting
the environmental impact of behavioural strategies followed by man and
thus advise scientifically for or against the use of a new technology.
Whatever form of government rules the future earth, they will be forced to recognise the need to live with nature and that all life, including humanity, is evolving, and quite rapidly at that. We can either take control of this evolution and guide it by working with nature, or let evolutionary forces destroy us if we destroy and do not respond to nature. Yet, in the end, man's innate yearning must turn away from base passions such as violence, hatred, greed and lust, towards God and a yearning for perpetuity in the hereafter under His guidance.
Back to Part I
Back to Part II
Go to next chapters: 1. ECOLOGICAL INTERACTIONS: MODELLING COMPATIBILITY.
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