INTRODUCTION

In the application of general system theory, a large effort has been to get at the systematic interconnections between systems ecology, on the one hand, and the evolutionary development of living systems and metasystems, upon the other hand.

Observations of the fossil record have yielded evidence of periodic mass extinctions, in which a large number of species periodically and suddenly died off and become extinct within a relatively short frame of time. In this process, we must picture the collapse of entire global or regional ecosystems that are no longer capable of supporting the kinds or degree of living systems that were previously supported. We must ask how entire taxa of animals, like the Dinosaurs, rise to ecological dominance for hundreds of millions of years, and then rapidly disappear from the earth with hardly a trace.

Taxon systems can be said to be large scale ecosystems that cover entire biomes, even regional, continental and trans-continental biomes, incorporating entire taxa of plants, animals, and decomposers. Taxon systems can be seen as the highest order integration of living systems short of the biosphere itself. We can for instance, in contemporary settings, compare North American biota with South American Biota, and with Eurasian or Asian or African Biota, and in each case one would find distinct regimes of flora and fauna that cross a numer of ecological zones.

Taxon systems reach deeply into natural time, and involve the cyclical rise and differentiation of many related species, generation upon generation, the flouressence of particular kinds of taxa in distinctive biological regimes. Species diverge and diversify when their adaptive success in a given setting is high.

We may thus describe and define taxon systems, largely the consequence of the rise of complex ecosystems based upon multi-cellular life forms that are sexually reproductive, as occurring simultaneously upon three systems levels:

1. The rise and fall of coherent reproductive communities of distinct subspecies or species in a given area or region and the processes of migration and dynamic succession associated with these patterns upon this level.

2. The rise and fall of branching families of related organisms, and their associated communities, including the intertwining of multiple families, over larger realms and biomes encompassing a diversity of ecological and environmental zones, along with the patterns of ecolocial shift, and succession associated with these patterns.

3. The rise and fall of large multiple orders of organisms in regional or interregional contexts, or continental or intercontinental contexts, and the associated patterns of shifting biome and community profile/diversity that can be classed as minor extinction events.

4. The rise and fall of entire phyla of organisms on a global scale that can be classified as a major mass extinction event and that can be associated with dramatic global shifts of environmental patterning.

We see in this scheme the encompassing of smaller taxon cycles within larger cycles, in which we can expect a high frequency and prevalence of level one cycles occurring, and relative rareness of major extinction events at level four. We can thus define a taxon cycle as the rise and fall of major branches and limbs of the family tree of life on earth, as evolutionary development in relation to ecological shift make possible growth and speciation of living systems in alternative and different directions. Niche expansion and diversification of a species or genus would be a signal of their adaptive success in certain regional biotic regimes. Niche reduction and loss of species diversity, perhaps connected to overspecialization, would signal the demise of not just a species, but an entire branch of that family tree, reaching back however many generations. If we find the loss of a certain species or kind of species in a given geographic area, we might be concerned about the corresponding lose of sympatrically related species in neighboring or similar areas. If we find losses across the board of this particular genus or family of species, then we can suspect global or regional factors impacting a range of zones and environments across the board. And if we find losses of entire superspecies in certain critical eco-niches at basic levels of the eco-trophic pyramid, then we should look for the impact of these losses at other levels of the pyramid and in other niches of the foodweb.

From an evolutionary perspective, eco-systems come to define a complex evolutionary framework of niches. I will call the "niche system" that comprise eco-systems and derive from the bio-geophysical background the evolutionary "super-niche." The essential characteristic of super-niches is that they are fundamentally social organizations, usually quite heterogenously composed of many different kinds of life that coexist in complex schemes of interaction. Evolutionary super-niches therefore set the common natural stages upon which the blind "forces" of selection are played out in the momentary lives of individual organisms. Eco-systems come to evolve in a coherent and systemic manner with their own cycles of succession, super-succession and climax. The term of evolutionary "saturation" and "supersaturation" describes the critical self-organizational aspects of evolving eco-systems that tend naturally toward increasing complexity at the cost of long-term stability. The dynamics of evolutionary process cannot be adequately explained outside of the framework of the coevolutionary eco-system. To deprive life of its self-organizing eco-systems is to force life into a pattern of extinction.

The point of departure for this chapter that I have entitled environmental fitness and adaptational selection, is to suggest that life creates its own contexts, and most life cannot occur outside of the biotic environments of which it is an intrinsically occurring part.

Mechanisms of population dynamics and speciation cannot be clearly understood or explained outside of such contexts. There is hardly ever a complete ecological vacuum in which a population can assert itself. No population runs its own history in complete isolation from other populations, and even if it did, theory suggests that eventually it would form its own subgroupings. It appears as if the primary mechanisms inducing extinction and selection do not derive intrinsically from the growth dynamics of a population itself, or necessarily from its speciational mechanics, but from the social contexts in which growth and development inevitably takes place.

Trait-fitness and trait-selection that are basic to the dynamics of populations and the mechanics of speciation are here construed in their social complementary forms as these are expressed within eco-systems. Environmental fitness therefore refers to a relative measure of "health" or "disease" within the environmental relations of the individual organism that would confer relative survival and reproductive advantage to it. Adaptational selection would refer to the patterns and abilities of other organisms around the population to successfully adapt to changing environmental conditions presented by the population in a manner that results in successful reproductive selection by that population.

Both concepts are rooted directly not so much in the trait-complexes of individuals or the trait-variability of populations, but in the functional behavior of individuals and range of deviation of behaviorial adaptations of populations that are an indirect consequence and natural outcome of differential expression of trait-complexes.

Furthermore, in being behavioral and functional in expression, both concepts point to a common form of fitness and selection that I will refer to as social fitness and social selection. In living systems, the significant behavior is social behavior. Thus, it is the indirect outcomes of social patterning of behavior, in terms of the complementary response patternings of other life forms to the actions and adaptations of the individual organism, that constitutes the basis of environmental fitness and adaptational selection.

Thus trait selection by itself does not explain speciation or evolutionary dynamics, unless we invoke social selection as its principal form of expression and determination in nature. This cannot be understood outside of the context in which selection normally occurs.

Social fitness and selection are in a sense primary pathways of behavioral and functional expression for natural trait-selection to occur. Speciation cannot be clearly understood outside of the contexts that relative patterns of eco-systemic social organization create. In other words, "traits" become "behavioral" through their social consequences, and natural "social relations" are defined through the behavioral consequences of trait configurations resulting in social selection.

In a fundamental sense, social fitness and selection are normal pathways that allow organisms as coherent interbreeding and feeding populations, at whatever point along the r-K continuum, to achieve and maintain a state of relative-K (r-K state). They represent the common strategic solutions that life can adopt to the basic challenges of the biological and evolutionary imperatives that are presented to them in the course of their lives.

Environmental fitness can be seen as the fitness that other organisms derive from and in turn confer to the actions of an individual organism in the expression of its own innate trait fitness. Adaptational selection can be seen more accurately as "counteradaptational selection," as the selection process impinging on other organisms that are the result of the adaptive responses of an individual organism to its own selectional constraints. These as a net consequence rebound upon the organism's own selectional patterning in a dynamic feedback process of mutual adaptational constraints.

To begin our digression, I will make the following basic statements about biological systems as natural information systems.

In keeping with systems theory, we refer to evolving systems of life as systems of energy transfer that accomplish a kind of work. The work it accomplishes in a basic sense is one of sustained life ("survival") and intergenerational transfer ("reproduction"). The work accomplished in the net result, is an increasingly differentiated system of life.

As energy systems, evolutionary systems are thermodynamic in character. This characterizes all living systems in distinct ways. All living systems have noise. There is mutational noise in the transcription and reproduction of DNA and in the loading of genetic variation in a population. There is ontogenetic and phenotypical noise in the variability of organismic development and functioning.

3. All evolutionary systems, as finite coherent systems, are in the long run subject to decay and death.

All systems, whether they are organisms or species or ecosystems, are subject to natural death. We know this to be true for individual organisms, and we know it to be intuitively true for species, but we must assume it is true especially for large and complex eco-systems. Observation of successional phases are the best examples of such eco-systemic death.

4. The basis for the "change" in biological systems is the replacement of the old and the dead by the new and the differentiated.

All systems eventually become replaced by new and different biological systems. It is the basis for all changes underlying evolutionary dynamics.

Thus we can say that the trend in evolution is to always go from simple to more complex states. This is the basis for a "progressive" nature of evolutionary patterning. The only way for such a system to be "simplified" or reduced is through death.

6. Evolutionary systems pull their resources from a common global resource pool, which at any one time is always limited in the total context.

At any one time, there is only a limited amount of the biological pie to go around, and all evolutionary systems share the same pie, such that what is taken by one system, can potentially be utilized by alternative systems.

7. Evolutionary systems eventually return their resources to the common global resource pool in order that they become available to new and different systems.

Death of evolutionary systems entails that the finite resources they utilized are returned to the common pool, and they stop utilizing energy resources that are then available to other alternative sysems.

8. In order for resources utilized by one system to be utilized by a new or different system, that first system must die or else change into the new system.

9. There is a net conservation of resources in the common global pool, though the form and distribution of these resources is changed through continuous patterns of death and replacement.

The continuous recycling of resources through the global resource pool by evolutionary systems in death and replacement leads to a net outcome of costs and gains of 0, such that there is net conservation of resources, inspite of local fluctuations of distribution and utilization patterns.

10. All evolutionary systems seek relative state equilibrium within the framework of the global resource pool by means of trying to maximize their share of the global resource pie.

In keeping with the principle of global conservation, evolutionary systems as working systems seek their own relative state of "equilibrium" as a form of "living conservation." We can translate this as biological "systems" trying to live as long as possible. They do this paradoxically, by means of attempting to maximize their share of the global resource pie. An evolutionary system would gain global unity with the global resource pool, if it came to encompass that entire pool within itself.

11. The contemporaneous coexistence of other, coevolutionary systems effectively prevent any system from achieving a condition of global state equilibrium.

No evolutionary system occurs in a complete vacuum of other similar systems. Therefore, on a very basic level, multiple systems are competing for larger shares of the global resource pie.

12. The total evolutionary system is constituted by the biosphere, and it encompasses all subevolutionary systems within its boundaries.

13. The biosphere is an evolutionary thermodynamic energy system. It is therefore entropic and in some measure of disequilibrium.

Just like the coevolutionary systems it encompasses and that compose it, the total biosphere is an imperfectly integrated evolutionary system that seeks global unity.

14. The biosphere never completely utilizes the total global resource pie, and thus obtains only relative state equilibrium.

Several kinds of conclusions can be drawn from this. Life as we know it will eventually come to an end as an energy system. As such a system, even if only one kind of life form could capitalize on the entire global resource pool, it would still obtain only relative state equilibrium. It predicts, as well, that a life form like Homo saipiens would eventually arise evolutionarily with the aim of monopolizing the total global resource pool for itself.

This basic thermodynamic paradigm describing evolutionary systems therefore embodies a fundamental feedback process by which old systems are replaced by new ones, and the new ones are both different, and increasingly differentiated, from the old ones. It explains the rise of gradational or stadial evolution that is seen as "progressive."

A model of this kind of paradigm follows. Imagine a single and supremely simple form of life that finds its way to a barren island absolutely devoid of any other life forms. This island is perfectly round and dotted along the cosst with many little tide-pools. Just above the tide pools are small streams that flow into the sea, and just beyond the streams are larger confluences of rivers that lead up to a central mountain, that is a volcano waiting to erupt. Conditions within the most proximate sea-side tide-pools on the island permit the survival of this single, simple life form. Because this life form is asexually reproductive, it rapidly mulitplies and fills in all the niches in a concentric band around the island, until its population reaches some carrying capacity of the outer ring of tide pools.

Because we are only visitors from another planet come to observe conditions, we take our notes and measures, leave a few markers, and then zoom away at light speed. Because we live for 10 billion years without growing old, we can return each million years or so to mark the changes that have occurred with our single life form.

In the first billion years, we notice few changes in our organism, except that now, instead of being only asexual, there are three varieties that are asexual, ambisexual and bisexual in its patterning. The ambisexual variety has sprouted little "roots" that allow it to remain attached to the sides of the tide-pools and survive conditions of drying and resubmersion. The bisexual variety has sprouted little tails that allow it to move to different corners of the pools.

In the second billion years, we see that both the ambisexual and bisexual varieties have come to invade the inner ring of tide pools, and the ambisexual ones have even moved onto the crevices of the spaces between the pools, and all in all we find 9 different kinds of life. The original assexual variety that just sort of floated at the bottom of the tide pools, until it happened to be washed between pools, spread out over a wider range, and invaded back into the sea where it occupied the range to a reef that surrounded the island.

If we continue our observations over the next two billion years, we will see some amazing changes happen to our simple life form, such that by 4 billion years, it has become a whole range of millions of life forms that inhabit the island at all levels. We find evidence that each .5 billion years the central volcano erupted, and slowly changed and added to the size the island.

We return one last time in five billion years to find the entire island extinct. The volcano that formed the central moutain apparently blew its top off, and all that remains is a huge lake surrounded by a concentric ring of coastline like an atoll.

We want to understand what happend since our last visit, and we dig down along the hills of the atoll to find remains of a highly advanced civilization, not unlike our own. We find some documents in a time capsule, and fortunately we have our universal decoding computer with us in our pocket. We discover that an intelligent form of life had emerged in the last half million years within the last quarter before our arrival, and they had designed a special anti-gravity reactor over the top of the volcano to create enough energy to drive an interplanetary space program. Their best scientists thought the volcano completely extinct, but they did not realize that a huge pool of magma rested deep down below a central vent system. Building the structures over the volcano effectively choked off the vent system, leading to a catastrophic explosion.

Thus seeing the end of all life on the planet, we decide to return to our space ship, and take one last look at a few tide pools at the edge of the ocean where it all began. We discover there through a microscope a simple life form floating around at the bottom of a couple of pools. We conclude our observations on a hopeful note that life will return to the island.

To make a long story short, if only one species found its way to a suitable but completely barren island on which a "ecological vacuum" existed, and if given enough evolutionary time, that species would go through a series of stages of evolutionary development, at the end of which would emerge a highly complex, multi-species system. Even if it had no social competition in the beginning, its own success and differential propagation would eventually lead it to create its own social competition leading to increasingly complex patterns of speciation.

These social evolutionary processes are by definition and invariably density-dependent relationships, and must be therefore construed from the standpoint of relative equilibrium about common hypothetical optima of "carrying capacity" that from an eco-systemic standpoint is referred to as "saturation" of the system. In contexts of "ecological vacuums" and of "supersaturation" it is expected that the density-dependent and therefore stabilizing nature of these relationships no longer hold, at least in a relativisitic sense. In such frameworks, things socially "fall apart."

An eco-system is usually defined by geophysical boundaries, even if such boundaries are behaviorally fuzzy and variable.

An eco-system therefore is a circumscribed area that has a finite if somewhat fluctuating set of physical limits, within which there is always a limited but variable amount of energy and biomass available to all those life forms that inhabit the region on a permanent or part-time basis.

An eco-system may be complexly organized in space such that it exhibits one or more core areas, an intermediate range of eco-clines, and a peripheral zone of overlap with other eco-systems that comprises the eco-tone. The structural relationships between these zones are important to the understanding of the dynamics of an ecosystem.

An eco-system can be measured in terms of its relative optima of equilibrium by how much the entire cycle of life is self-contained within its bio-geophysical parameters. It is something like a terrarium in which continuous evapo-transpiration within the bottle recycles water to the roots of the plants. Closed feedback loops contained within an ecosystem define the density-dependent parameters of the system.

No eco-system is completely bounded. All eco-systems are partially interconnected to the global biosphere. Interconnections of the ecosystem to the external biosphere are a source of disequilibrium to the system, and are the basis for the introduction of density-independent factors to the system.

Eco-systems are by definition natural social systems within which different interacting forms of life carve up the pie-of life and occupy a range of niches made available by adaptation to the variable contexts and settings that contain eco-systems. All interactions between organisms are de facto social and involve some level of "mutualism" and "competition" at the same time.

An eco-system composed of even one kind of organism would still constitute a social eco-system if a population of such organisms were contained within a semi-closed environmental context that is somehow self-sustaining. In a sense, a petri-dish with a slime mold is a very rudimentary eco-system, at least for the life of the mold.

This suggests that even the very first primordial and "prototypical" life forms had to have arisen in an environmental context that was somehow minimally eco-systemic. Through its own self-replication this life form would have created its own life-context, and hence, the basis for its own evolutionary development.

Thus, consideration of the evolutinary implications of eco-systems has an important bearing on our equations of fitness and selection, as in a sense it inverts these values such that they are no longer considered the outcome of individual adaptational responses to conditional mechanisms imposed from without in complex environmental settings. Instead, these settings themselves become to some extent the product of an organism's behavior in its natural world.

At the same time, it is evident that the environment comes to instigate itself into the life-world of the organism, into the organism itself, in complex ways.

On the level of the ecosystem, this problem comes to be defined in terms of the following questions. How does an ecosystem composed of interacting species within a food-web or life-pyramid come to create "boundaries" about itself that serves to maintain the stable long-term equilibrium of such a system? While individuals, groups and even entire species may come and go from such systems, the system remains, for a period of time, relatively stable and consistent in its patterning of relations and populations.

Within such systems, most interacting species arrive at or develop common "solutions" that are coevolutionary and mutually interdependent. They become mutually trait-adapted to their contexts, and this sense of adaptation serves to hinder intrusion from and prevent loss to the outside world.

It can be seen that a form of adaptational selectionism comes into being that is fundamentally social, intra- and interspecific, and follows a kind of "strategic channeling" of adaptations toward a common direction. It creates a boundary between those species within the system that are trait adapted, and those outside of the system who may want to invade at some level. It leads to competitive exclusion and inclusion. It also tends to make it difficult for member populations to simply leave the system without suffering severe deprivation and external competititon. Selection on this level might be thought of as competitive and behavioral in nature.

From the standpoint of the average individual of the group, that organism has come to define for itself more or less a clearcut niche where many periodic interrelations with other kinds of organisms are to be expected. Reactions and interrelations in such contexts tend also to be fairly well worked out.

Fitness from this standpoint becomes fundamentally "counteradaptational" and coevolutionary. It would be the measure of the individual's capacity to respond effectively to the subtle perturbations of relationships that might occur in such systems.

In such contexts, saturation would be reached where species gradually switch from being opportunistic in reproductive patterning towards greater stability and equilibrium. This transition would be reflected in many aspects of their life, and would be reflected in changing profile of their population parameters.

Such systems would exhibit patterns of balancing and stabilizing selection, in various stages of their development. At the same time, in the later stage of their development, they will become increasingly susceptible to disruptive factors of selection that are either density dependent or density independent.

What is needed to be defined is a complex cycle of adaptation and fitness at this level. It is a derivation of the taxon cycle, but interspecifically oriented.

A one-to-one correspondence between a gene and a trait is a naive and oversimplistic view of how genotype comes to affect phenotype. Traits cohere as trait complexes, such that mutations may on average influence outcomes across a suite of traits, rather than at any single point. Most traits are either polymorphic or pleiotrophic, or a complex combination of both. Genetic organization is undoubtedly complex.

Genetic solutions to the problems of survival and reproduction were arrived at only through billions of years of evolutionary experimentation. Each individual organism represents a special experiment of evolution. Will the organism survive and achieve reproductive success with its given suite of genetic traits? It is becoming increasingly apparent that genes act in combination to produce sophisticated transformational algorithms in the cellular differentiation of the body. Suites of traits that affect the total organism are produced in trait-complexes. These trait complexes integrate to define the individual as a unique member of a species--a characteristic phenotype.

When selection occurs at this level, it is my contention that selection is operating in relation to such trait-complexes, and not on single or "point" trait characteristics alone.

First, random genetic mutations may have a greater chance of influencing some aspect, or multiple aspects, of such a complex, or of multiple integrated complexes, than if that mutation is tied exclusively to a single genotypical trait.

Secondly, random genetic mutations, unless they proved extremely deleterious, would likely have little net outcome for the total organism, either positive or negative, except that it might nudge that individual a little further or closer towards the evolutionary goal of achieving adaptive fitness. Most point trait-changes can therefore be considered relatively null from the evolutionary view of the long run, and large populations can normally support a huge genetic load without failing.

This suggests that genetic complexes, as worked out evolutionarily, are on average relatively stable though they are also basically flexible and malleable. It sets fairly broad tolerance limits to the genotypical and phenotypical profile of the population as a whole, allowing a broad range of variation to be normally exhibited.

This also suggests that as evolutionary experiments, genotypes of most species are heterogeneous and are in a sense "predisposed" as organizational complexes towards achieving adaptive fitness with the environment.

Even in strongly opportunistic, r-selected species that regularly suffer high percentages of mortality, any species has worked out a solution for the problem that, under normal circumstances which can subsume a broad range of environmental variability, assures the survival of enough individuals in each generation to achieve not only population replacement, but growth.

I would claim that, short of the complex brain, this is the primary mechanism that life has utilized for "beating" the odds in the game of reproductive survival, and it is this kind of mechanism that accounts for the alleged natural intelligence of biotic systems in general. On another level, it is a kind of "problem solving" that has led to a sense of gradational and stadial "progressive" evolution.

I will distinguish four main levels of biological patterning to deal with in a systems approach. These levels are:

Micro-systems approach, which involves the organiismic functioning of an individual of a species

Species systems models, involving intra-specific and inter-specific relations between organisms

Global life systems models, which include the entire tree of life, and the current evolving biosphere.

I will attempt to explicate each level for the insight it might derive for both biological systems and for systems theory. Many aspects of these systems are fairly well understood and much better elaborated than can be done here. This systems approach is only a vehicle to provide an alternative means for thinking about the inherent complexity and variability of life on different levels of its occurrence, and the vast network of interdependencies that it represents.

Especially, I wish to emphasize the holistic and integrative aspects of biological systems upon each level, and between the three levels, such that these systems are to be seen as super-critically interdependent and contextally self-organizing. They create their own contexts for understanding, and have thus led to their own form of natural adaptive intelligence, of which we human beings are end products.

Group adaptational mechanisms may be inherently more flexible and hence evolutionarily variable than for even the individual itself, and may confer to different groups possibilities of evolutionary development that wouldn't otherwise exist. Group adaptations can foster stable conditions leading to sustained positive selection in a larger context.

Adaptational selection can be referred to as any behavioral adaptation of an organism or a group of related individuals that changes the outcomes of the evolutionary imperative for other organisms or groups within a well-defined ecosystem.

Biology is about life, and the exact scientific definition of life is a complex one. Basically, any living system must be a self-replicating or reproducing system, and must be self-sustaining for its cycle of existence within some environmental framework. For all life on earth, the enduring and universal characteristic is that, no matter what form or pattern that life form assumes, it has a characteristic DNA genetic structure. It is the continuous replication and modification of this structure and the development of systems of survival by life forms to accomplish this genetic transmission, that defines biological information patterns as we know them to occur on earth.

The genetic structure of life on earth is not necessarily obligatory by any basic physical principles. It is entirely possible to imagine an entirely different kind of genetic structure from an alien life form, and this is likely to be what we find.

All the necessary informational patterning necessary for life on earth, for its replication and survival, is found in the genetic encoding of the DNA at a molecular biochemical level. DNA contains the genetic totipotency of all biological informational patterning on earth, and constitutes the foundation for biological information systems.

Furthermore, biological systems develop and evolve. They are living systems, and therefore they have a natural life cycle. They are born, go through some kind of growth and maturation process, and eventually die. By contrast, physical systems are by definition dead or non-living systems. Finally, all biological systems evolve, or are evolutionarily dynamic, in the sense that they reproduce themselves in an historical series of generations, and are thus transformed into new and different systems.

It is my contention that physical systems, though possibly developmental with their own sense of physical history, are essentially non-evolutionary systems in the same way that we construe biological evolution to take place on earth.

I believe it is impossible to imagine a living organism that is completely and absolutely separate from or isolated from other living organisms. For biological life, as far as we know it, exists within a larger living system of other life forms, and this has been true probably from the very first emergence of life on earth.

I believe that DNA alone is necessary but not sufficient informational patterning for biological systems to occur and survive and to reproduce themselves. First, as noted above, all instances of DNA structure take place in the context of populations, and usually of populations of different kinds and forms of life. Population genetics takes on a more complex dynamic than simple genetic encoding alone, and frequently has to do with the relative patterned occurrence of alternative traits or mutations of the DNA sequence that leads to increased repoductivity or otherwise.

Up to this point, I've said little that is new or that is not better understood by most biologists. Beyond the level of population genetics, there is another level of systems theory in biological research that is in general referred to as "ecology" or "eco-systems" approaches. Eco-systems concern the informational patterning that is in the environmental relations established by the living members of a species in a "niche" or within a larger framework, that must be mastered by that species in order to achieve survival and reproductive success.

To go one step up the ladder, so to speak, I would suggest an even higher level of informational patterning that is occurring within biological systems, and that I believe to be necessary to the survival and success of species within these systems. I would call this level that of regional and global biological regimes, which contain complex patterns of information that structure the frameworks of survival for many, if not most, species within its framework. In this model, few biological systems are totally isolated from other systems. If a system is cut-off from a global system, it is likely that it will reach a kind of evolutionary cul-de-sac in the development of its life forms.

In its simplest form, a biological regime can be considered to be a kind of biological supersystem in which there are numerous different species functioning at multiple levels and in differing areas, that are necessary for the continuance of the system as a whole. A great deal of life, in fact, most if not all of it, has been critically shaped within the totalizing contexts of such macro-scopic biological regimes.

Species present a specific reproductive boundary that cannot be crossed. Yet at no point in time, can we exactly identify "speciation" as an event that serves to permanently isolate two species from one. Only long-term isolation or relative behavioral segregation of two populations of a single species, resulting in different adaptational strategies and in different genetic profiles arising, can account for speciation in a stable way. The challenge is that though speciation is known to be a result of extended periods of time, at no point in time can we clearly distinguish the reproductive separation of a single species into two except in the form of reproductive isolation that occurs after the fact of speciation itself.

There is a relativity of macro-evolutionary regimes also. The species and adaptive suites at arise in the context of any particularly epoch are not necessarily transferrable or the same as those existing for any other epoch. This is especially true when ages are compared that are separated by relatively long hiatus or lacunae of time.

It is quite apparent that not all selection that occurs in an epoch is necessarily natural nor "adaptive" from the standpoint of selecting for a more fit species. Frequently, rates of death far exceed rates of replacement, or else, in a surge of population expansion in the rapid exploitation of some open niche, it is possible that individuals reproduce without necessarily being the most fit for survival. Survival of a species or a particular population seems far more complex and prone to chance events than something like the law of natural selection alone would fully account for.

Any number of fairly random or chance factors can come into play in determining the survival or extinction of an individual, a group, a population or an entire species, much less the simultaneous or concurrent extinction of multiple species. To identify a prime mover or unicausal explanation for any complex set of events represented by mass extinction, especially as this unfolds over time, is to underrate and oversimplify the complexity of the connections surrounding such events.

Living systems are complex systems that, in total, create their own environments indirectly in a web of life set of interrelationships. Biotic environments are vast and usually all encompassing. Species of life generally move and exist within the frameworks of these biotic environments, and not outside of them. At this macro-systems level, all life is a complex patterning of interaction and interrelationship within which there are few recognizable physical boundaries. Of course, zebras normally do not cross the Atlantic by themselves to populate the wildlands of Wyoming, unless someone brought them in for the purpose. But there is a sense that the physical ranges of plants and animals in different regions overlap and interlock in an unending pattern that carries itself into other regions and areas of the earth.

In such a regime, most effects are complex and are the indirect results of other patterns. In fact, from the standpoint of natural systems theory, biological systems demonstrate an inherent complexity at almost every level of their manifestation that is at once beautiful and intricate.

Little attention to date has been paid to such a level of biological information patterning. It is my interest in this chapter to focus mainly on this, and upon the consequences and understanding of the basic issues of what I consider to be the current biological regime compared to previous regimes on earth.

Extinction is a natural outcome of evolution, much as death is a natural consequence of life.

Mass extinction as the natural outcome of supersystems and evolutionary aegis or regimes.

Biological supersystems constitute entire evolutionary epochs during which a particular predominant adaptive regime or aegis occurs.

Evolution has been considered to be "blind" and this has been somewhat dogmatically defended by its faithful followers who see anything otherwise as being either the hint of Creationism or else the cutting communist hand of Lamarkianism. Thus, genotypic and phenotypic traits are strictly dichotomized, and natural selection always proceeds by chance and circumstance, about as blindly and unbiased as statistics can become.

But models of alternative intelligence suggest that this apparent "blindness" may in fact be only a kind of inherent "myopia" of life that has been self-organizing, but informationally complex and extremely synergistic. Evolution may proceed therefore in a manner that is not exactly or totally "random," but has a "non-random" directive component about it, especially in its more advanced stages, that suggest an incipient natural form of primitive "quasi-intelligence." Intelligence can be considered in this sense a natural derivative of the complex self-organization of natural information systems. Even human intelligence is organically derived as the by-product of millions of years of convoluted evolutionary history.

Life is totally related. We all derived from the same single proto-biological source. In other words, we must speculate that the real "Eve" of life was a primitive prokaryotic germ in some kind of complex chemical soup. In fact, spontaneous generation of proto-life forms may have been multiple and relatively independent, and may have resulted in the overlapping of two or more different fundamental forms, and perhaps the extinction of some original forms and their lineages. But for all intents and purposes, a single source will be considered. This means at some level very far removed, we are cousins of amoeba, sequoias, giant kelp, snails, squids and the bacteria that infect us and make us ill.

The tree model of life is from the standpoint of systems design is very informative at all levels.

The DNA can be considered more than just a blue print. It is a set of working transform operators that allow a complex scheduling of cellular differentiation to take place all more or less timed to a genetic developmental "clock" that is in essence the result of the number of genetic transcriptions and replications that occur. Exactly how this genetic clock works remains mostly unexplained.

We can picture any DNA, at the moment of its conception as a fertilized cell, beginning a process of serial growth, cellular replication and differentiation until an organism of specific description is reproduced. This pattern of development can be described as a tree.

All DNA clockwork takes place within the environment of a cell. A cell can be understood as a kind of minimal unit of life defining the biological environment of the DNA it contains. This cell interacts with the outside environment, usually with other kinds of cells. Growth and development of an organism can be understood in terms of the increasing pattern of multiple replication and organized differentiation of its cells, in the construction of a complete and mature organism. The model of a tree diagram can describe this for any organism. Each tree would look differently for each organism and for each kind of organism.

The cell is the natural and self-organizing environment of the DNA. DNA cannot survive and replicate outside of this environment. Even modern genetic engineering has not succeeded yet in creating organisms from test-tubes. They can manipulate the DNA in cells, and even extract and implant it in different cells, and even grow cells in different matrices, but they cannot undue the environment of the cell itself.

It is my contention that all life creates to some extent its own minimal environment, at whatever level, and that in the total sense the entire biosphere becomes the global environment for all Life. This self-organizing environment is a principle pattern of intelligent adaptation of life, within which evolutionary development achieves some measure of non-random direction that we associate with primitive intelligence.

From this standpoint, evolution proceeds primarily from the standpoint of the successive generational replication and populational multiplication of these "tree structures," with a variable of change introduced such that there occurs slight variations between structures. No two structures are exactly identical. Variation of structure reaches a point over time, especially when external selective factors are introduced, that essentially two or more kinds of tree structures are produced that form essentially different and non-interbreeding populations.

Thus given enough time, the tree structure of a single organism turns into a plethora of alternative tree structures of different kinds of organisms. That organisms of one generation die, to be replaced by the next, means that we end up in time only with the results of evolution, and feint records of its natural history. The larger tree structure can only then be inferred by indirect evidence.

Eventually, some of the tree structures sprout legs and eyes and ears, and grow brains. The tree structure is a homologous model of evolutionary development that applies on two levels simultaneously. It applies in terms of the cellular development of DNA in the growth and development of a single organism. It applies in terms of the tree structure of the differentiation and development of different species of organisms in relation to one another. In between these two levels of tree structures, is a vast forest of trees of all kinds of living creature, all interacting together. Each individual organiismic tree structure is related to all other organiismic tree structures on some level or another.

The larger level of evolutonary tree structure is a model that is only to be inferred from the evidence of past and present generational instances of trees, and from the interrelationships among the trees of the present and past forest of life. It is in a sense an abstract and eidectic model, whereas in theory at least, the organiismic tree structure of an individual organism is held to be actually quite mechanical, concrete and substantive.

This opens the door to a kind of detailed systematic analysis that replicates to some measure the complexity of life we construe. This kind of analysis would be based on a complex data-based connected to a kind of inference engine. There would be several levels of nested discrimination tables within this database, all interlinked to one another, such that changing values in one area of the database, reverberates in alterations of values throughout the entire database. Each discrimination table would comprise the total calculus of variables at each level and in each area, going from the organiismic level, to a populational--interspecific level, up to a global level.

In this regard, informational patterning of Life, and the inferrable self-organizing proto-intelligence that its evolution suggests, occurs at four basic levels of informational stratification as natural systems.

First, we seek to understand how processes of organiismic development occurs in terms of its DNA transcription, replication, cellular differentiation and resulting cycles of growth, maturation and eventual death.

Second, we seek to understand the dynamic patternings of the functioning of the organism as a whole, and especially how it survives in the world, in its total life-context, and manages to reproduce itself to continue the chain of life.

Third, we seek to understand how populations of such organisms function, especially in relation to one another, reproduce and managed in the process to change into multiple forms.

Fourth, we seek to reconstruct the entire tree of life, and also to understand how the dynamic functioning of the interacting species of living forms occurs and affect changes upon this tree in any given period or place.

In the first level, I will speculate that "adaptive intelligence'" evinces itself in terms of the variability that is inherent to organiismic development and growth. The core driving mechanism of evolutionary development and speciation is held to be the process of genetic mutation underlying selection and drift of entire populations. This process is held to be entirely by chance occurrence. Indeed, it is a random process, but there is a sense in which the cellular environment, and by extension the extracellular environment, begins to have a shaping influence upon the pattern of development almost from the moment of conception.

If certain damaging chemicals are present in the early stages, dramatic deformations of the finished form are predictable. Of course, this does not directly affect the genetic structure itself, unless it does do this--for instance by means of radiation or chemical alteration of the DNA by some outside influence. Damage or variation of the cellular structure surrounding the DNA will not alter the DNA, but will change the ratio of survivability of the organism as a whole, and thus its likelihood of being replicated. Up until this point, evolution at this level is essentially blind and random. God has played dice with Life.

The environment intrudes upon the epiphenomenal patterning of the genetic development of an organism from the moment of its conception until the day of its death. This does not usually directly affect the genetic structure of the organism except by chance mutation. But the net result is always the same, changing the chances of survival and reproduction of the organism. This is seen as a fundamentally blind evolutionary process, but it is also the case that it opens the door a little wider for a kind of adaptationism of the organism that may in the final analysis not be so blind. That the body of organisms can adapt themselves to changes in the environment can be seen in several ways--homeothermic mechanisms that maintain internal body temperature, instinctual and reflexive behaviors, or the utilization of anti-bodies & white corpuscles to attack foreign disease elements in the body. These are all, of course, products of evolutionary development, i.e, genetic adaptationism.

At the second level, success of the individual organism is defined primarily in a larger context of its social grouping within which it is situated. This in a larger sense is situated in a context of environmental relations and relations with other kinds of social groupings and other kinds of beings. This complex context ultimately manifests itself in the everyday life-world of the individual organism, and the pattern and capability of the organism's behavior and responses within such contexts have a decisive impact upon the organism's chances of survival and reproducibility.

Often, events can occur in other contexts, only indirectly related to the context of this individual organism's life world, that can have a critical shaping influence of that organism's chances of survival. A volcanic eruption half a world a way can precipitate a climate change pronounced enough to inhibit growth patterns of certain plants, altering the relative availability to an organism of certain kinds of plant foods. Similarly, a surging population of some remote species could cause migration patterns into an area, as a secondary effect, of another kind of species, resulting in loss of food and habitat for an organism.

Most species are socially defined in terms of their reproductive patterns, such that in such contexts individuals cannot be isolated from the context of their own or similar groupings and hope to survive. A butterfly migrating in North America may be blown off its course by a storm and find its way in Britain, only to discover that it is effectively isolated from its own kind so that it cannot breed and contribute to its "gene pool" in a successful way. Thus it dies not fulfilling its evolutionary purpose though it may have been a wonderful survivor.

At this level, there may be many indirect causes that influence the individual organism's chances for survival and reproduction, and this suggests a broader model for "natural selection" than is normally implied by this term. The conventional prejudice of natural selection is the image of a lion hunting a hyena, or wolves scavenging a herd of bison for the sick and young. It is an image of active predation and competition between different species in the same contexts, leading to limited survivorship--"survival of the fittest."

That natural selection itself may be much more of a numbers and odds game than any one imagined, and that its forces may be more strongly counteracted by adaptive mechanisms of the individual, often on the spur of the moment, suggests that the game of life is more complex and less controllable or well-defined than people would like to have it.

Humans are discovering how complex the indirect chains of life may really be when they find DDT levels going up the food chain, or that the massive kills of fish in the sea may be indirectly the result of too much fertilizer leaching off the lands. This complexity demonstrates the interconnectivity of the entire web of life, such that its difficult to find many examples of relatively, much less totally, isolated species or even small ecosystems of a handful of interacting species.

Few boundaries on earth are naturally or completely impenetrable by all forms of life, and life tends to adapt itself to suit a very broad range of different kind of habitats, even very unusual ones like the sulphurous waters of hot springs, geo-thermal vents or even the boiling water of geysers themselves.

It is difficult amidst such complex interconnections to tell simply or clearly where decisive or significant determining factors may lie, and how different sets of factors may interact in even more complex ways to alter the chances of survival for an organism. More often than not, determining factors may be beyond the control of an organism, like the surge of carbon dioxide from the bottom of a volcanic lake that wipes out entire villages, or the eruptions of a volcano that destroys all the life upon its slopes, or a forest fire that catches many species in its swath of destruction. In such dramatic and catastrophic events, there is little individuals can do to adapt themselves and survive. Disease epidemics in the New World and Old are clear examples of this. People die off in massive numbers from forces they do not even understand or see, much less know how to adapt to. Seldom can they run fast and far enough away to escape them, before it catches up with them and consumes them along with most of the others.

I believe that just as a DNA is wrapped within a nucleus within a biotic environment of a cell, individual organisms of species tend to wrap themselves in enivironments that I will call their life-world habitats. These habitats usually include other individuals like themselves who cohabitate the same area, and who often interbreed and share other support mechanisms. There is a level of intraspecific mutualism that helps to hedge the bets for an individual organism's survival. Maintaining a habitat, and at times imposing such a habitat upon the world, helps individuals to beat the odds. Thus even for normally migratory animals, it can be found that they have a home range that they have chosen for the purposes of setting up nests and breeding grounds.

The notion of habitat addresses the life-world of the individual organism in a holistic sense, and must be capable of providing on a daily or weekly basis the required resources necessary for that organism's survival. Thus, the total habitat for a particular organism would be complementary to that individual's daily life world experiences and requirements for survival. Habitat looked at from this way is a kind of common "pathway" or set of pathways an individual can choose to take from one event to another in the course of a day or a week or a month.

To some extent, an individual will "explore" its life-world and create a habitat, mostly based on some ingrained "template" of experience. This process of exploration will open the door for "intelligent" adaptation of a specific individual or specific group of interrelated individuals.

Thus, normally, a population of such individuals or groups will be dispersed out over a wider range of territory with very similar modes of adaptation. Beavers may find places to build their homes and dams in lakes, streams, ponds and rivers. Especially in many life forms, it seems as if the young of the species are deliberately "sent" on explorations of their world to establish themselves in some kind of niche within the larger framework.

By such means, individuals gain a handle of control over their life-worlds that guarantees a certain order and predictability about it, and that hedges their bets against not surviving in a world that is otherwise stacked against them. Coming to "know" such a habitat and to successfully adapt within such a habitat, to the point of being able to successfully reproduce, represents a major accomplishment in the life -world of any organism. They are driven to this task instinctively. Gaining such knowledge and success, organisms gain an inherent advantage and measure of control over their life-worlds, such that those organisms that intrude upon it will be in an inherent disadvantage.

On this level, it is not just a matter of individual survival that we are talking about, but of group survival. Individuals cannot survive long on their own or away from the context of their primary social group. In this regard, there is strength in numbers. Few catastrophic events are so destructive that they are capable of destroying entire groups, though it sometimes happens that way. Even if most people were destroyed by some lethal disease chances would remain strong that a minority can usually survive to propagate and reproduce.

The reproductive mechanism at this level of the social group is a social mechanism. The group, especially an organized one, confers survivability upon its members that its members wouldn't otherwise have. A lone cow away from a herd would soon be victim to some predator. Its chances of successfully reproducing are far better within the herd than outside. Some kinds of species, especially predatorial ones, for instance, snakes, actually spread out into wide areas as individuals, and only reaggregate for the purposes of procreation. Such creatures cannot support themselves when their densities reach a proportion that they are competing among themselves for the same sources of food. But even such creatures must remain within some "home" range of the larger grouping, such that they can periodically reproduce themselves.

From the standpoint of population genetics, populations of interbreeding individuals generally "radiate" out across a suite of ecological niches, with small groups of such individuals finding for themselves habitats across a broader range. Invariably, this can lead to some adaptational flexibility that leads to active exploration of the environment. Adaptive radiation is a sign of reproductive success of a species that has gained the upper hand in adaptive survival. Individuals are able to push beyond the boundaries of the preestablished population to extend the ranges of adaptation of the group as a whole.

At this stage, standard continent-island and bottleneck models of gene flow become appropriate to the description of basic processes of speciation. Large populations may come to occupy very large ranges that incorporate a wide variety of different niches. Subsequent events may result in the relative isolation of subgroups from the main host, or a fracturing of the main body into separate subgroups. If effective isolation is maintained long enough, then definite speciation and reproductive boundaries between the subgroups occur.

Adaptive radiation of a species into large areas enters the game of adaptive survival to another regional and interregional level of information. At this stage, it is the interaction between different kinds of species that comes to the foreground, and at this level, the conventional "competitive" model of natural selection seems most appropriate. Interaction between species is not always directly competitive, and can frequently become mutualistic or else parasitic. At this level, different varieties and kinds of life forms are exploring complex adaptational spaces and relations, and creating viable eco-systems that confer some adaptive advantage for individuals at a higher level of biological function.

At this level, individuals are working not mechanically as a member of a larger, single population, but organically in differentiated roles within interdependent niches. Organisms carve out niches within larger zones, sometimes in overlapping bio-zones, and come to be interdependent with the niches and functioning of other organisms in complementary ways. Even predation and competition can have net positive effects on the predated species if it leads to the culling of the weak and the diseased and the maintenance of optimal population levels.

At this level, adaptationism frequently comes at the intersection of distinctive zones, and comes to embrace the entire tree of life on some level. Micro-organisms, plants, mammals, insects, fish, bacteria, algae and fungi all come to co-reside and help maintain a vast and finally organized biological system of interspecific function. The ocean is a good example of how vast such a system can be, as it presents few natural barriers to movement, and thus consists of a vast region of overlapping ranges and habitats for a very wide variety of life forms.

At the highest level, we can consider the interregional aspects of adaptationism of all of life itself. Carbon, water, nitrogen, oxygen, the basic components of biological life forms, all have their natural cycles in the world, and different species in different zones and regions affect and impact upon these cycles at different tropic levels. It is at this level that we have become most concerned with the long-term effects of human selectionism and socio-environmental circumscription. Over fishing in many regions of the world has resulted in major drops in fish populations globally, to the point of no return for many species that were once abundant and plentiful. The loss of a few species can spell disaster for other species that depend upon them in some indirect manner.

At the core of eco-systems evolution is a model of social relations that can be applied differentially to most if not all forms of life interactions within such systems. The basic model derived herein is based on competition theory that has been well worked out.

Before undertaking an explication of competition theory, it is important to note several caveats:

1. All natural social interactions within eco-systems are by definition both competitive and mutualistic at the same time. To the extent that eco-systems are closed affairs with finite enegy economics and biomass productivity, any organism represents a percentage of the total. Multiple organisms therefore exist in a fundamental way at the possible expense of other alternative organisms. This is fundamental natural competition that can even be applied to highly social creatures that exhibit traits of altruism and kin-selection. Thus, we must conclude that the gene is, in the most fundamental of senses, naturally selfish.

At the same time, all social relations within eco-systems are also to some extent inherently mutualistic in the sense that by fact of social interaction any set of organisms are contributing to the overall maintenance of equilibrium within the entire system. Lions may take down large prey, and this is perhaps the ultimate of competition, "survival of the fittest," but the Lion could not exist outside of the framework of the prey it hunts and depends upon. Even extremely asymmetrical social relationships are therefore, in the total scheme of things, mutually interdependent relationships. This speaks to the degree of functional integration of the entire system as such. We all have our parts to play in the natural scheme of things, and each organism, in its time, plays its own parts.

It is true that all systems are not perfectly integrated, and therefore there is much room for error and correction to be made in such systems. "Noise" can be introduced into the "normal" functional patterning of interrelationships in a number of ways, leading to destructive consequences within the system. Destructive consequences occur not when there is no more competition or no more mutualism of interdependencies within the system, but when environmental factors sweep through in independent fashion and impact on all organisms in an "equal" manner. It is when Lions go out to hunt, but find nothing to kill, or, alternatively, there are no more Lions to hunt the game that usually expects to be hunted by Lions.

Such relationships, both normal and abnormal, ordered and noisy, speak of the fundamental density-dependent nature of social relations. I would like to suggest the following basic paradigm:

With increasing saturation of social relations within a biotic system, there is a move along a density-dependent continuum such that species move from a mode of fundamental "inclusive fitness" towards a condition of increasing "exclusive fitness" that leads to interactive patterns that are more complexily organized. These tend to emphasize the survival and reproductivity of the individual (individual fitness and selection) over that of the group (group fitness and selection).

In saturated models, the principle of competitive exclusion in its most extreme forms can be assumed to be of strongest value.

This fundamental mechanism serves to promote greater relative-K for any reproductively coherent group, up until a level of saturation of the eco-system is reached. Saturation I will define as the level of ecosystemic-K or what I will call "social-K." There is a long-term natural tendency for such systems to grow gradually out of balance, by means of counteradaptive bioschismogenesis between interacting groups or populations. The natural outcome is for supersaturation as a social, density-dependent condition, to accumulate in the core of such systems, leading to the consequence that relative exclusive fitness, defined as an increasingly competitive model, no longer enhances systemic stability, but begins inducing negative, destructive feedback.

At this stage, death rates increase comprehensively as birth rates remain low. The system, as a spiraling thing, grows out of balance until it begins, like a spinning top, wobbling out of control.

Thus we can imagine a continuum between "inclusively fit" social patterns and "exclusively fit" social patterns along which all organisms within such a system range. These are the social interactional correlates of "r" and "K" in basic populational and speciational models. I will call them the relative social fitness of any organism or population within an ecosystem.

2. All ecosystems, being by definition social systems, are partially open to the larger global ecosystem framework. This means that all patterns occuring within an ecosystem are relative to that system as part of a larger system.

Overpopulation or high-density values of such a system are in essence forms of "local overpopulation" in a larger context of relationships. If a population experiences a condition of local overpopulatin, a common and direct solution is migration out of the system to a larger context. The result of such migration is usually dramatic reduction of population attendant upon widespread broadcast dispersal or increased risk of negative selection due to increasing uncertainty factors.

In general, it can be said that the "exogenous relationships" maintained from without the system serve as density-independent factors that impinge upon and influence the system as independent variables of change.

Likewise, it can be asserted that "endogenous relationships" tend to be maintained as stabilizing and density-dependent factors and occur as dependent variables of change and variation within the system.

This gives to any organism, and any set of organisms, or group or population, within an ecosystem, an intrinsically "dual" identity both within the system and without the system. Species boundaries usually criss-cross multiple ecosystemic boundaries in complex ways. It means that usually a single species will at least potentially inhabit and function in multiple ecosystems, and possibly at multiple levels within different ecosystems.

It is therefore the case that individual organisms and groups can function in ways within ecosystems that are in response to relationships that are occuring outside of the relative parameters of an ecosystem. In other words, there is room for "confusion" of roles between endogenous and exogenous relationships between organisms. If a migrating herd of buffalo was annihilated in one region, then their ability to return to another region where lions can hunt them may be fundamentally impeded.

3. Applying formulations from 1 and 2, the possibility exists (in fact quite commonly) that for any organism or species, there are differential pressures endogenously and exogenously exerted that promote both inclusive and exclusive patterns of fitness within an ecosystem, leading to a differential oscillation of that population between alternative modalities. Thus a population that exhibits relative-K or social K within an ecosystemic framework may be driven by external factors independent of the system to revert to a more non-K state, throwing the internal system into a state of disequilibrium.

In this context, it is possible to imagine a case of overpopulation leading to a splitting of the population into two groups, driven by the differentials of adaptation between the internal and external frameworks.

4. To complicate this picture one step more, if a species or population inhabits more than one ecosystemic framework at one time, patterns driving social-K and social selection between these different frameworks may lead a population down different pathways fundamentally. This is of course commonly observed in nature. What achieves relative K in one adaptive framework, may not be what allows a subpopulation of the same species to achieve relative K in another framework. This will lead to a bipolar or multi-polar divergent pattern of social selection of species within alternative ecosystemic frameworks.

This brings up the question as to whether the relative social integration within any ecosystem can achieve such influence or power to influence the speciational patterns of a breeding population. Ample evidence of sympatric speciation exist without needing to invoke actual isolation of populations. The core of the ecosystem exhibits a kind of evolutionary gravitational pull on surrounding species to its center.

To some extent, this gravitational pull can be seen as merely a function of distance itself within an allocation of energy budgeting. This suggests a kind of central-place model of ecosystems. Species that have to invest a great deal of energy migrating between different eco-systems must trade-ff energy that could be invested in either survival or reproductive success. In fact, distance to ecosystems is what morphological size is to populations, and the key trait of m-selection and selection of morphological independence, are important components of the ability of species to inhabit and occupy multiple ecosystems.

Thus, it can be stated that for any given species in any given niche, there is an optimal geographical limit or boundary, a "zone" beyond which the loss in energy required to migrate, coupled with the increase "risk" involved in distant migration, would outweigh the gains to be achieved from increasing one's resource base.

But it can be seen that gravitational "pull" can be exhibited even in complex settings where absolute distance does not seem to be a critical factor. In such conditions, density dependent relations of saturated systems would entail that the likelihood of gaining a position of greater equilibrium outside of a system will be less than the loss involved in losing one's niche within the system.

So great is the inherent drive for equilibrium that is a natural outcome of the evolutionary imperative, that it can split a single population into two, creating a social-functional boundary inbetween where none existed before. In its initial stages at least this does not have to be defined by any inherent differences of trait configuration. It may arise exclusively as the result of different selectional patterns operating on different modes of behavioral and social functioning within the different ecological zones.

It is known also that inbetween such stable ecosystems, "ecotones" exist that characterize an intermediate transition zone between these systems. Boundaries divide regions between ecosystem cores as "eco-clines" that mark a gradient of transition zone between the cores. In such "edge" systems, as saddle nodes in an evolutionary landscape, and as "neutral" or "no organism's land," it is my hypothesis that relative K is difficult to achieve and sustain, and that such zones are characterized by high rates of disequilibrium by selectional patterns characteristic of isolation and peripheralization.

Such eco-clines and ecotones are usually depicted geographically in vertical and horizontal terms. In this regard, I wish to emphasize the importance of the "intermediary" zone, that first 100 feet above and below sea-level incorporating coastlines, lower river systems and lakes, as one of the most important "global ecotones" that exist where characteristic patterns of ecotones can be studied in abundance. In fact, this global ecotone, as an intermediary zone between aquatic and terrestrial zones, constitutes one of the most basic divisions found in life between ecosystems. We may characterize all ecosystems as either terrestrial, aquatic or intermediary. From an evolutionary standpoint, I believe that the intermediary zone has been the most biologically dynamic and evolutionarily productive set of ecosystems that has ever existed. The intermediary zone also exhibits all the features of succession.

These considerations naturally lead us to an understanding of "competition" as a basic model of natural social relations. Before proceeding, it must be reemphasized that "competition" can be construed in different ways from different points of view. Counteradaptation and coevolutionary speciation can be considered a form of competition, as can sexual selection and social strategies of reproduction. Parasitism and predation represent basic forms of competition, as does heliotrophism and the competition for sunlight by plants. Schools of fish may school not because they are minimizing their own chances of predation, so much as they may be attempting to maximize their own competitive advantage over their sibling organisms, or over other schools of fish.

I would say that underlying all forms of competition is a form of systemic relational interdependency, such that we can speculate on the following kind of formula:

Greater density-dependency of a relative social K-state is correlated with greater social interdependency between organisms and populations, and this is manifested as some form of social competition between organisms, often mediated by social structural patterns.

Whatever form of competition becomes expressed, it leads to an emphasis of exclusive fitness of individuals over groups.

Greater exclusive fitness begets greater adaptive fitness of individuals in an ecosystemic context, leading to greater reproductive success of the individual within the context, even at the expense of the reproductive success of the population to which the individual belongs. Even if drone honey bees are by evolutionary law destined not to contribute to the species gene pool, (exhibiting marked "inclusive fitness") it is certainly also the case that a few princess bees compete ruthlessly for the exclusive advantage of being the queen and sole contributer to the next generation of bees.

Competition theory is derived directly from the logistical formulas relating to density-dependency and shares its basic assumptions, which are known to be unrealistic when applied to actual populational parameters and conditions. These are known as the Lotka-Volterra Equations. Consider any two groups in competition N₁ and N₂ with respective carrying capacities of K₁ and K₂ and their own maximal instantaneous rates of reproduction r₁ and r₂. The simultaneous growth of both groups co-occuring is given bya pair of differential logistic equations:

Where a in both formulas is a competition coefficient which measures for each group its competitive inhibition per individual on the other group. Competition coefficients are normally numbers less than 1.

In the absence of any intergroup competition, the competition coefficient and the N factor of the other group in each equation equals zero, both populations would grow sigmoidally according to the Verhulst-Pearl Logistic equation, attaining independently their carrying capacity.

The inhibitory effect of each individual in its own group is by definition 1/K of that group. The inhibitory effect of each individual on the other group is given by the respective competition coefficient of the other group over the K of the other group, or a/K. Outcomes of competition between the two groups depends on the relative values of a and K for each interacting group, such that there are four possible sets of outcomes depending upon these values:

To understand these possible outcomes, it must be asked at what density of group 1 individuals will group 2 individuals be held in check (at zero growth) and vice versa, such that the density of each will prevent the increase of density of the other. When the density of one group equals the inhibitory effect of the other group upon it, K₂/ a₂₁ or K₁/ a₁₂, respectively, the density of the other group cannot be changed.

In a competitive vacuum of the other group, each group would increase to the limit of its carrying capacity K, and decrease above that limit. But in the presence of K₁/ a₁₂ individuals in competition, N₂, N₁ decreases at every density, and vice versa.

While in the logistical formula for a single population, r decreases linearly with increasing N until reaching K at which it is zero, in the competition equations, there is a set of lines relating r to K in each group. Each line corresponds to the differential population density of the competing group.

If the values of the equations above are set to 0, then we can derive the boundary equations for increase and decrease for each of the populations, such that:

If these two linear formulas are plotted on the same set of axis, the isoclines dN/dt for each group are given below which each group increases and above which they decrease. These lines represent equilibrium population densities or "saturation" values of the ecosystem. If the combined densities of both groups lie above the line, neither group can increase. If the combined densities lie below the line, both groups can increase.

In consideration of the four sets of possible outcomes, only one set of outcomes leads to stable equilibrium between the groups, when neither group is able to achieve densities that are greater than the other. Implicit to this inequality between groups allowing for mutual coexistence is a condition such that "each group must inhibit its own growth more than that of the other species." (Pianka 178) If carrying capacities of the two groups are unequal (as is usually the case) then stable equilibrium can still be achieved if the product of the two competition coefficients is less than one. If population sizes of both groups are below their respective carrying capacities, niether population achieves the potential carrying capacity that it would if it were not in competition.

By multiplying through the equations with r1N1 or r2N2 respectively, we can yield equivalent equations that express the competitive variables more clearly:

Where z is equal to r/K for each subscripted group and ß is za for each group respectively and where the first term to the right of the equal sign represents the density-independent rate of population increase. The second term measures the intraspecific self-dampening rate and the third term the interspecific competitive inhibition of the rate of increase.

These formulas can be written for a community of n-species in the following way:

At a steady state, at which the value of the equation is 0 for all groups i, the equilibrium population densities are given by:

In this equation, the larger the final term to the right of the equation becomes, the more competitors any one group has, and the more distant that groups equilibrium population size (relative K) is from its independent K value in a competitive vacuum. This is referred to as "diffuse competition" and refers to the total competitive effect of the remainder of the community on a particular population.

The equations have been extended to partially embrace a model of predation, which can be considered a specialized form of competition.

Where the first equation stands for the prey population and the second for the predator group. These equations are solved by setting them equal to 0 and factoring out N to get the actual rate of increase and then setting this value to 0. There are not self-limiting density effects as in the competition equation such as -zN², but each population is constrained by the other. In the absence of any predators, the prey population would increase exponentially without limit. The number of contacts between the two groups are the product of the densities of the two species N₁N₂. This value is multiplied by a constant p₂ to represent the maximum rate of increase of the predator population. Multiplied by p₁ the term appears as a negative value and represents the corresponding decrease in the rate of prey population.

The prey population reaches equilibrium when the predator's density equals r₁ /p₁and the predator population reaches equilibrium when the prey populations reaches d₂ /p₂.

Each group's isocline corresponds to a discrete density of the other group such that below a prey threshold density, predators decrease, and above that threshold density, they increase. If the predator's density increases above a threshold, prey density decreases, and vice versa. Though joint equilibrium exists for the two groups, the densities of both groups do not converge upon this point. Any given initial values instead result in predictable predator-prey oscillations of certain magnitudes. If the two groups are near the point of joint equilibrium these oscillations will be of low amplitude. The solution to this formula is therefore periodic, with the cyclical changing of group densities and increasing disequilibrium developing over time. These conditions are termed "neutrally stable" and are generally unrealistic since most populations have either self-regulating or density-dependent inhibitory mechanisms.

If a self-dampening term -(zN²) is applied to the equations, joint equilibrium is reached and there is dampening in the oscillations. A realistic self-dampening term for the predator should include a prey-density dependent factor, yielding the following equations:

The prey equation is the simple competition equation but the predator equation is a function of the relative densities of predator and prey. The predator population is fundamentally dependent on the prey population and cannot increase unless the prey population exists. This equation is held to be unrealistic in conditions where there are more prey than the predator population can exploit, such that the predator population cannot be directly the consequence of prey densities without some threshold effect occuring.

These equations also don't take into account differential response patterns by predators to increasing prey densities, termed functional and numerical responses. Predators have a limit of satiation beyond which they change their rate of predation. Also, increasing prey means increasing predators, increasing the rate of predation. Complex systems models taking these types of factors into account have been developed, but are not elegant for generalizations. If carrying capacities of prey are assumed relative to carrying capacities of predators, such that there is equilibrium of the prey population, and the carrying capacity of the predator population is assumed to be relatively independent of that of the prey, influenced by outside competition, then a complex quadrant model is yielded such that:

In this model, differential magnitudes of changes in population densities of both groups determine the stability of equilibrium such that resulting vectors spiral inward (damped oscillations) toward the intersection of joint equilibrium (where both equations are set to zero.) Else the vector spirals outward (increasing disequilibrium between the two populations), or else they spiral in a circle about the point of joint equilibrium (neutral stability). These kinds of oscillations suggest population "cycles."

It suggests that if predators are inefficient, neutral stability will result. If the predators are very efficient at feeding, then disequilibrium will eventually result until a threshold is reached when there are too few prey that the predator can no longer efficiently exploit the prey population. The predation drives itself to increasing inefficiency, requiring greater efficiency until a minimum threshold creates maximum ineffiency. The result of the last instance of an overly efficient predator would be the predator driving itself to extinction along with its prey. In this picture, damped oscillations would result from some omptimum threshold of predatory efficiency, either by counter-adaptational patterns of the prey that serve to limit predatory efficiency, or else competition by other predators and the presence of multiple kinds of prey packages. Dampening oscillations are the most frequently observed in nature, while disequilibrium cycles are rarely observed. Dampening oscillations suggest mutual equilibrium being approache as the densities of predator and prey track one another towards greater stability.

It is assumed that more efficient predators that can increase their rate of reproduction at lower prey densities will replace less efficient predators, and this will in the long run serve to reduce the stability of the system. Balancing counteradaptations of prey that can more efficiently escape increased predator efficiency should restore balance to the system. If this does not happen, then an efficient predator can be expected to drive both populations to extinction. Perhaps T-Rex was the most efficient predator of all, and slow moving, cumbersome supersaurs reached the zenith of their K-development. In such a hyperdeveloped system, this would be an expected outcome.

Symbiosis is another derivative of the basic competition equations by simply changing the signs of the alphas to positive and altering K's to X's since they do not represent maximal population densities, hence:

Equilibrium conditions can be represented by a pair of linear equations such that each population reaches equilibrium at density X in the absence of the other group, and increasing the density of one group results in increasing the equilibrium density of the other group. If both densities are positive and the alphas intersect, then stable joint equilibrium exists at a point of intersection of the two isoclines. The following quadratic table represents the alternative conditions then possible:

Again, these assume ideal conditions presumed in the original competition equations. Conditions in which one population is considered as neutrally independent, while the other derives benefit, might also be safely modeled as a combination of the predator and the symbiotic equations together, borrowing assumptions from both that relate to such a model.

Of course, these equations derived from the Lotka-Volterra competition equations only hold under unrealistic conditions where the carrying capacities, rates of increase and competition coefficients are assumed to be constant and the common environments the groups inhabit are thought to be homogenous and there is no time lag between changes in density. Inhibitory relationships between groups are always linear, and each individual of one group is considered equal to a corresponding individual of the other group. It is true that these values vary with population density and other factors intrinsic to different kinds of life forms. There is always some lag between changes in density, and environments vary contantly in place and time.

Though these equations are unrealistic in their assumptions, and lead to complexity problems in the enumeration of their variables, they have been very productive in providing a conceptual framework for understanding competition between groups.

It is clear that these equations always presume conditions of equilibrium between populations, but in fact these conditions may frequently not exist in ecosystems. Unsaturated conditions would entail that mutual dampening interactions between the two groups might become density independent as conditions of disequilibrium are maximized. If a system goes to saturation between two competing groups, usually one group is driven out or to extinction by competitive exclusion. Differential natural rates of increase and relative carrying capacities between groups usually entails intrinsic inequality between the two groups such that one group will reach saturation before the other which is still increasing. This leads to the elimination and competitive exclusion of the saturated group, by the fact that its population declines after its rate of increase reaches zero.

The principle of competitive exclusion suggests that two groups with the same ecologies (two predators at the same trophic level and in the same niche) cannot coexist in the same time and place, except that one will be driven out or to extinction. The corollary has been that if two groups coexist, then there must be ecological differences between them (two different groups of competitors occupying different trophic niches). While in its extreme form this hypothesis is regarded as untestable and unrealistic, conflating patterns of individual variation, it does emphasize that some ecological difference is necessary for mutual coexistence of competing communities in saturated environments.

The principle of competitive exclusion gives us a handle on how most species are driven either to extinction or towards marginalization.

So far, the principle of competition applies, as in its extreme form, to two different groups. It does not apply in exactly the same way to the consideration of intraspecific competition by individuals within one group, which often leads to opposite consequences. In general, intraspecific competition is held to affect the population's tolerance, its increased use of broad-based resources and wider phenotypic variability. Whereas interspecific competition is held to generally restrict a group's range of resources and habitats, intraspecfic competition is held to lead to results that expand or diversify or increase the range of resources and habitat.

Given any resource pool or bounded habitat, the intraspecific population is expected to spread itself out unevenly across a gradient or continuum that represents differential utilization patterns. This results in a distributional patterning within an ecosystem for the same group, such that competition between members varies along the gradient, and such that there should be differential levels of individual fitness. It is expected that individuals should behave in order to equalize the ratio of demand to supply along the gradient, equalizing the intraspecific competition.

This notion assumes that lower intensities of competition between members of a group will be negatively correlated with higher fitness values for these individuals. It is not clear that this is the case in highly competitive populations, especially in contexts of breeding competition. Competitive exclusion within a group still tends to be a factor, I believe, in the core region of a resource zone where competition is greatest, hence density-dependent factors play a greater role.

Formulas for intraspecific competition may be adapted directly from formulas for interspecific competition, if we assume that populations can be subdivided along lines of clinal variation represented by their fitnesses and relative densities into two subpopulations that are competitive with one another. Resulting clinal variations suggest the ecoclinal gradient from the core areas of ecosystems. The result is internal stratification of the ecosystem of a population in a manner similar to the stratification between two populations.

In the long run, increasing intraspecific competition will lead evolutionarily to selection patterns that result in congenic interspecific competition between closely related species.

Patterning of intraspecific competition must be itself a relatively density-dependent phenomena, such that with increasing densities, increasing exclusive fitness sets in. It is the case that narrow margins of relative-K for a group in competition with another group, will lower the threshold at which competitive exclusion begins working within a group as well, to the point, in extreme conditions of supersaturation, it becomes a 'war of all against all."

It is held that intraspecific competition can lead to dispersion and or expansion of a population, such that the net result is an increase in the variety of resources and habitats used by a population in less saturated peripheral zones. It can also lead to an increase in the trait-variability of the individuals themselves that are better adapted to marginal conditions. This only holds when marginal zones are assumed not to be saturated.

In this context, intraspecific competition has different selectional effects on groups than does interspecific competition between groups, the latter being restrictive and inhibitive, the former being diversifying and expansive.

It is argued that in general at equilibrium total intraspecific competition has the net effect of balancing total interspecific competition. If this is the case, then it can be assumed that increasing intraspecific competition should lead to increasing interspecific competition until saturation is achieved. It also suggests that groups with higher levels of intraspecific competition should be capable of resisting increasing levels of interspecific competition from other groups.

In fact, it appears to be the case that conditions of high interspecific competition can result in patterns of relatively low intraspecific competition within each of the groups, as a result of cooperative social organization by the groups in the face of their common enemy.

In other words, in the face of mutual exclusive competition by other out-groups, members of an in-group can be expected to adopt patterns that lead to greater relative "inclusive fitness" by members of the group, through social organization of the group that serves common goals of maximizing relative fitness and minimizing negative selection.

Such social organization does not usually entail "blanket equality" of all members, though it has been argued that it leads to a relative "equality of fitness opportunity" of all members through clinal distribution. This is especially true for r-type life forms. But social organization is more often as not accomplished through stratification of the group, such that some members are favored at the expense of others. Such stratification implies internalized competition that has been structurally regulated (relative social-K) for the benefit of some at the expense of others (kin-selection).

Conditions of saturation are held to induce increased competition and competitive fitness. Many patterns are associated with increased intraspecific competition, including: delayed reproduction, smaller clutch size, larger size of offspring, parental care, mating systems, dispersed spacing systems, territoriality. It also is held to account for ecological diversification leading to niche separation. This is the kind of speciational patterning that reflects diversifying and balancing selection, and reflects increasing intraspecific differentiation of a species within a stable saturated ecosystem. It is alleged to lead to optimization of utilization of minimum determining resources.

As a result, it can be assumed that increasing competition and saturation of eco-systems leads towards either extinction, competitive exclusion or to increasing differentiation within the system resulting in attainment of stable relative social-K states.

Complex systems models of social selection are derived from an understanding of social interactionism within shared contexts. These models are tied back to the basic aspects of the model of differential trait-fitness and selection considered in previous chapters, and it is demonstrated that processes of selection and fitness that drive evolution cannot be understood in a strict cause and effect framework. The problem of fitting evolution into a causal framework is really a hen or egg dilemma. To see evolutionary development in terms of the speciation of a single population outside of changing social contexts is to attempt to explain evolutionary processes in linear terms.

Only by construing evolutionary dynamics from the standpoint of recurring social cycles within larger natural cycles can we derive a more accurate systemic model of evolutionary proces. These cycles may lead down different developmental pathways, whose various stages have expectable consequences within an information systems framework. Only in this way can we resolve this kind of hen or egg dilemma that has been at the background of the understanding of natural selection from the beginning.

Models of cyclical process that reflect the fundamental and general realities of evolutionary development can be built. The model I propose is that of a periodic oscillator. Any energy system that is bound to a stable state of equilibrium, such as a fully saturated ecosystem in a range of fairly stable environmental parameters, by some "restoring" or self-regulating force, which I take to be mechanisms of social selection based on reproductive competition, will upon disturbance from its equilibrium position, "resonate" at a frequency established by the reproductive rates and death rates of the populations involved. Achieved relative equilibrium of any population is a measure of its "evolutionary inertia."

This oscillation tends to be driven periodically by a complex set of external forces that impinge upon the system in expectable intervals derived from the oscillation patterns of neighboring ecosystems.

The preceding digression based on theories of competition demonstrates several things. In general, increasing competition between forms of life tend to lead to a pattern of exclusion, such that other kinds of relational values are excluded between such life forms. We can say that in general, as things tend toward relative K, things also tend toward increasing competition. In the extreme form of competition, total exclusion results in either extinction or marginalization.

Relational interactions that do not reflect direct competition, can be considered inherently and indirectly competitive, but are to be seen as efforts to maintain relative equilibrium in conditions that would otherwise result in disequilibrium or exclusion.

Thus complex social organization and patterns of counteradaptational selection and coevolutionary interdependence arise precisely in conditions where potential competition can be expected to otherwise intensify. There would be no need for social organization or for complex patterns of interdependency to arise in conditions where there is no competition as a result of saturation and relative K-states.

Thus it can be seen that competition constitutes a basic mechanism governing and leading to trait-displacement in natural selection and patterns of speciation.

Social interactions between and within groups in ecosystems tend towards increasing complexity and are difficult to generally model in realistic terms. Nevertheless, it is evident that most forms of interaction can be at least partially depicted through competition, which illustrates a basic principle. Given any two (or more) organisms (or groups) in a finite resource system, a basic density-dependent relationship is inherently established, such that increasing growth will result in competitive constraints operating between all coexisting populations. Complex patterns of symbiotic mutualism and social interaction are derivative consequences of these basic constraints. While this model describes mutual coexistence and the rise and declines of populations about some hypothesized state of optimal equilibrium, they do not describe the resulting patterns of social selection that can be expected from them.

Exclusive fitness and direct social competition are positively correlated with density-dependency and relative saturation within a system.

With increasing saturation of any system, it can be expected that social selection will manifest itself in increased rates of premature (nonreproductive) death and dampened actual instantaneous rates of birth.

In highly saturated, competitive environments, some species will increase at the expense of others that will face either extinction or marginalization.

Any system must eventually become unstable if some species cannot be displaced by exclusion from the system, or the system cannot achieve a higher threshold of equilibrium.

Unstable systems will result in relative innate competition that is density independent in its function, returning the entire system through increased death rates to a lower level of saturation. We may say that a form of nondifferential negative selection sets into the system.

This suggests that there is an inherent long-term instability of all ecosystems that will tend eventually towards disequilibrium in spite of relative states of achieved mutual equilibrium between members of the system.

We will go back to our basic formulas, and demonstrate that any presuppositions of density-dependence results in two-way interactions between any two organisms, groups, populations or species. The following kind of "interdependency" paradigm hold generally true for any kind of social interaction we may wish to represent in time or place:

A + B	B gains + 1	B neutral 0	B loses -1
A gains +1	Both gain	B 0, A + 1	B-1, A+ 1
A neutral 0	B+ 1, A 0	B 0 , A 0	B -1, A 0
A loses - 1	B+ 1, A-1	B 0, A -1	Both lose

I will call this framework a discrimination table of basic interdependencies. We may hypothesize that any interaction, or any predictable set of similar interactions, between any set of individuals, groups or populations, regardless of the specificity or inequality of the compared terms, can be placed in one of the sets of squares, and in one square only. The same interaction cannot be placed in two different squares at the same time. Thus, the absolute value of the table as a whole will be equal to total number of finite interactions or relationships recordable, within a given area over a given period of time. This might be called the functional density of an area that would be a measure of the relative density-dependency of that area as well as of the relative saturation of the area and indirectly a measure of species diversity and heterogeneity.

We would of course add cells to the table in a third dimension if we which to specify relations occurring between three or more compared terms and can be represented on an enlarged squared table. The range of possible interactions can be specified for any number of terms, as well as the degrees of freedom.

This table is called a table of interdepedencies because it presumes a basic principle of density-interdependence operating between any two or more organisms, groups, etc., within any finite system.

1. It is the natural imperative of each represented group to maximize its share of resources within an ecosystem. (innate competitiveness hypothesis)

2. Each represented group will strive to minimize its loses within the ecosystem.

3. In the growth of such systems, it can be expected that eventually the gain of some will come at the expense of others.

4. Direct competition should emerge as the result of increasing densities of populations and net saturation of the system.

The center value where interactions are "mutually neutral" would in an absolute sense be nonexistent or incorrect, if we assume a basic assumption of innate competition. But in a relative sense it is very possible to describe the mutual coexistence of different life forms that have no direct consequence upon one another. Innate competition is probably under most circumstances a residual and negligible factor in fitness and selection patterns, unless a case can be made for total supersaturation of the area in question. At the stage where innate competition would become a factor, it can be assumed that it becomes indirectly a density-independent factor, as it would probably affect all organisms in the system in the same proportionate degree. There are many contexts in which different species are not only mutually tolerant of one another, but actually indirectly codependent upon one another.

We can say therefore that relationships tend to move away from the center of neutrality in one or another direction. We can say that maximum ideal equilibrium would be achieved in the upper left-hand corner of the table, and maximum disequilibrium in the lower right-hand corner. It will be demonstrated that probably both states are never achievable, and therefore most social relationships range between the two extremes.

Before proceeding with our model, it is necessary to emphasize the concepts of relative ecological rarefaction or saturation of an ecosystem. These concepts of rarefaction and saturation are related to the notions of carrying capacity, equilibrium, density-dependency and climax within a region, but they point to the energy-dynamics and bio-geo-physical resources of the system, especially as these are stratified between tropic levels. In general, saturation of any area can be considered to be the relative degree to which the total energy budget and biological resources of any system, and therefore biomass productivity, is used up by the life forms existing within that area. A saturated system is therefore one that approaches the maximum limit of the system's total carrying capacity. A rarefied system is one that approximates some minimal level of resource utilization within the system.

The concepts of saturation and rarefaction lead to consideration of heterogeneity and species diversity found within such systems and to a complex table of allocation of systems resources distributed between different kinds of coexisting life forms. The increase of resource utilization by one lifeform in a system will lead to offsets in the levels of utilization by other lifeforms.

This table of complex resource allocation within any eco-system I will call the functional trophic-taxonomic matrix that underlies the functional dynamics of the system. Any system compries a range of niche potential at multiple trophic levels, and becomes representative in time of a variety of different kinds of organisms that seek to inhabit various niches at different levels.

In general, the table would look like this, and is derived from the matrix and pie-of-life model developed previously:

Consideration of "neutral" relationships invokes models of matrices and life-pies previously described about the basic pattern of relationship occuring across Kingdoms in any ecosystem. In this model, relationships occuring across the basic divisions of Kingdoms present some of the most fundamental differences that can occur between organisms sharing a common environment.

This model suggests a basic functional stability of relationships tending towards what I will call minimal r-equilibrium (or maximum r-disequilibrium) found in all ecosystems, and that underlies the evolutionary stability of the entire biosphere. The stability of all nature rests on the fundamental interdependencies that arise on this level of interaction between different primary trophic levels. At this level, competition can be expected to be minimized. Minimum r-equilibrium would represent the minimum threshold of adaptation for a population. This minimum stability underlying all ecosystems occurs at a threshold of maximum rarefaction that a system can achieve and still remain a coherent system. Thus, we cannot in reality ever presume a total or perfect ecological vacuum occuring.

At the same time, this same model sets upper limits of K for all the primary trophic orders within the total system, such that changing equilibria in one of the orders must affect the other orders somehow. This upper limit defines the upper threshold of adaptive K-equilibrium for any population.

This would also set finite limits to total carrying capacities of any one primary trophic order, as well as a sense of resonance fluctuation of trophic limits within each order and between orders that describes a cyclical feedback pattern that can be either dampening or amplifying in nature.

It is understood that all organisms share density-independent values of innate competition, and consumers share a fundamental dependency upon producers. It is possible to imagine a browser that grazes itself to extinction if it is specialized on one kind of plant, while producers indirectly depend upon both consumers and decomposers. Most models of direct competition are at this level specific to the Kingdom being represented. We expect certain forms of competition between animals, especially at the same trophic levels that we do not expect between plants and animals. Plants also compete typically with one another for sunlight and other basic resources.

Furthermore, it is the upper levels of the pyramid of life where we expect to find the greatest amounts of direct competition between species, that we conventionally stereotype as "survival of the fittest." We also expect to find the greatest amounts of direct competition within trophic levels rather than between trophic levels, though it is understandable that there is significant competition between trophic levels, especially those that are contiguous with one another on the pyramid of life. There are few hard and fast rules in this modeling of social interactions between different kinds of life, because diversity of species and interaction is the rule rather than the exception. Complex food chains and cyclical systems develop within the pyramid of life such that many kinds of indirect relations are established.

In order to get a handle on the meaning of innate competition in its varying forms, it is necessary to distinguish between innate types of interdependent competition occurring along some kind of competition continuum that includes all possible interactions between organisms.

I will hypothesize that competition can be seen in two basic forms that are related to the selection outcomes they favor or result in. These two forms of competititon I will call reproductive competition (r-competition) and adaptational competition (K-competition). I will state that r-competition leads towards reproductive success or failure of one organism or set of related organisms, in relation to that of another. I will state that adaptational competition (K-competition) leads to adaptational success or failure of one organism or set of related organisms in relation to that of another.

If we go back to our original formulas, we can see that K-competition is an independent variable in the biological imperative, and that r-competition is a dependent variable.

In any given interaction, we can always assume some minimal level of K-competition occurring between the agencies, but we do not have to assume r-competition occurring except under certain conditions.

Whereever we find r-competition occurring, we can expect some degree of K-competition also to be occuring on a more fundamental level.

In general, I will state that the more different organisms or set of organisms are in both functional and taxonomic patterns, the greater the degree of adaptational competition can be expected between them and the less the reproductive competition. Vice versa, the more similar two organisms or related set of organisms are to one another, the greater will be the degree of reproductive competition between them.

Adaptational competition can be construed as encompassing a broader spectrum of interaction in which interactions between agencies or parties do not necessarily alter reproductive fitness values of either group, but alter the adaptive fitness values of the group.

In a sense adaptational competition sets absolute limits to the carrying capacity of any unique or related grouping, compared to other groupings that are different from it, beyond which relatively density-dependent limiting factors become relatively density-independent factors. K-equilibrium is the natural expected outcome of K-competition, and is easier to establish between species that are widely divergent on the trophic-taxonomic matrix than between those that are closely related.

What exactly distinguishes reproductive r-competition from adaptational K-competition is the issue of relative exclusive fitness that serves to emphasize the selective exclusion of the individual compared to that of the entire group. In a sense, therefore, K-competition compared to r-competition is just the social interactional inversion of our notions of r-K fitness and selection values. R-competition leads to greater relative K-fitness and selection, and results from this patterning. K-competition results from and leads to greater non-K or r-fitness and selection patterns that can be said to be characterized by inclusive fitness.

It can be expected therefore that r-competition results in equilibrium between and within related species whereas K-competition tends to lead to adaptive disequilibrium between related species. K-competition only leads to equilibrium as a function of the "evolutionary distance" between the interacting species.

We can claim that reproductive-competition results in reproductive-selection which tends to narrow the intrinsic trait-variability within a population by means of exclusion and emphasis on exclusive fitness. Reproductive competition is therefore a death-instigated selection process that leads to greater r for one group while maintaining K for another group.

On the other hand, adaptational-competition would result in adaptational selection and counteradaptational selection that would tend to broaden the intrinsic trait-variability represented by a population by means of inclusion and emphasis on inclusive fitness. Successful adaptive competition is birth-instigated selective process that should result in increasing reproductive rates leading to K.

To understand this, we must seek to understand the idea of indirect social selection, and how forms of competitive exclusion can actually result in greater equilibrium and balance between different species. This is accomplished by ecosystemic compartmentalization, or the separation and reproductive isolation of similar species, where all naturally occurring systems would tend, through natural increase, towards disequilibrium anyways.

In other words, we cannot hypothesize an innate mechanism within a species that would automatically tell it to curtail its reproductive rate under conditions near equilibrium. This is in spite of increasing death rate that should offset the rate of birth at and beyond equilibrium as this is expressed in carrying capacity or relative saturation.

The presupposition in the basic population and competition formulas is that there is some internal "balancing" mechanism in the organism or population especially, that says to it "slow down reproduction" once conditions approach optimum. In general, death rates and birth rates are only indirectly interdependent. Not only is there an inherent lag and differential distribution of instances of deaths and births over time but the classical equilibrium formulas imply a causal interdependency that doesn't necessarily exist.

At the stage of equilibrium, some other set of mechanisms must begin to kick in to "regulate" the cycle between deaths and births. These mechanisms are not directly the density-dependent factors of stress and strain on basic adaptational resources that result in increasing rates of death. Nor are they mechanisms like predation or coadaptation that counterbalance or offset preestablished reproduction rates.

They are mechanisms that arise intraspecifically and congenically, and in niche competition between functionally similar kinds of species, They result in the competitive separation and reproductive isolation of subgroups or organisms between the two populations. It leads to clinal distribution and divergent speciation even in sympatric contexts where no physical barriers are seen to exist.

They always are most marked in conditions aproximating relative-K between the organisms or groups involved, when the resources profiles they share are the most similar and therefore the most strained. At this stage, either organism or group, in order to increase its reproductive rate, must do so at the exclusion of the other group. The group cannot otherwise continue to grow.

The obverse side of this is to consider the basic adaptational competition between to widely divergent forms of life, such that the common overlap in resource profiles between them is very narrow. Such species can tolerate high mutual densities of one another without requiring competitive exclusion.

Adaptational K-competition becomes most marked in conditions between divergent species when there is some minimal resource (or set of resources) shared between them in a profile such that density-dependency of relationship arises in what would be otherwise relatively density-independent contexts. It indirectly affects the rates of reproduction and death between the two groups. This can arise from conditions of environmental fluctuation. Other wise, it would be most marked in contexts between trophic levels such as strong predation or extreme parasitism, when the existence of one species comes to depend exclusively upon and utilize the other species as the principle and only basis for its resource. The rates of reproduction of the predatory or parasitic species or group drive the other species or group into extinction or marginalization.

In this sense, competitive exclusion can result from extreme forms of either strong r-competition or K-competition, which suggests that the most evolutionarily stable patterns are intermediate between the two extreme "strong" forms.

Reproductive competition can therefore be seen as a special form of adaptational competition that occurs when two groups greatly overlap in their resource profiles on the trophic-taxonomic matrix and are competing for reproductive advantage, or r-fitness, between one another within a shared context. It may also arise when the results of such competition are expressed in terms of relative r-fitness values between the two groups. It can occur between life-forms that are not directly intra-specific, as for instance congenic sibling species, though reproductive competition at the intraspecific level is expected to be the greatest, leading either to organismic spacing, territoriality or complex forms of social organization.

It can be seen that both kinds of hypothesized competition are in fact interrelated to one another, such that adaptational competition leads indirectly to reproductive competition, and reproductive competition is always fundamentally a form of adaptational competition.

We can say, paradoxically, that reproductive competition always leads to interspecific patterns of exclusive fitness, whereas adaptational competition always encompasses the entire range of relative fitness values, whether it is exclusive or not.

Comparison of adaptational and reproductive competition supports the following kind of representation:

Considering this framework, we can hypothesize the following kind of generalization:

At any functional level of trophic-taxonomic classification, we can distinguish between inter-group and intra-group forms of competition. We can hypothesize that at any level there is a characteristic degree and type of adaptational competition occuring between representatives of different groups.

The closer the groups are related in both taxonomic and functional identities, the greater will be the direct reproductive competition between them. Another way of stating this is that the degree of trait-overlap or similarity on the trophic-taxonomic matrix, between any two or more comparable organisms, groups of organisms or species, the greater the inferrable interdependencies between them are likely to be expressed in terms of exclusive fitness and reproductive competition.

In such contexts where very similar kinds of life come into interaction, the net result of such interaction must eventuate in some form of relative isolation or mutual exclusion between the two forms. Succession of biotic forms in certain regimes can be understood as a consequence of this operational principle. Fundamental differentiation of speciational processes underlying all evolutionary processes can be understood in this way. The result of this patterning is also to set up a variegated topography of isoclinal zonations in the distributional patterning of different forms of life.

Organisms that are sufficiently divergent from one another on the trophic-taxonomic matrix, and in which mostly adaptational competition occurs, can mutually coexist within the same habitats and environments without this adaptation leading to mutual exclusion or the creation of functional boundaries between the populations.

Complex patterns can result where it is possible for adaptational competition between two widely divergent forms of life to result in changes in reproductive competition for either form of life with some other closely related forms of life.

Social selection operating on interdependent populations must be construed from the standpoint of the long-term evolutionary consequences of such systems. Obviously, systems that drive towards extreme mutual disequilibrium reach natural limits of maximum rarefaction of interaction at which point inclusive fitness and r are maximized and density-dependence is of minimum value. In a sense, differential selection increases as the rate of reproduction increases and inclusive fitness kicks in, such that it is in the conditions of relative disquilibrium that we find the fastest rates of evolutionary development. This model is depicted below.

In such a model, it is evident that a changing rate of evolutionary development for any given line, in any given ecosystem, must be an inverse function of relative density-dependent relationships, such that increasing states of disequilibrium result in increasing rates of speciation. It would suggest that there is a fundamental lag time in this process, which is the equivalent to the lag time between birth and death rates in normal populational dynamics. I also hypothesize that there is a line at which optimal selection values occur such that selection processes occuring above this line have fundamentally different consequences than selection processes occuring below this line.

We may make a distinction between differentiating and nondifferential selection processes, whether they occur above or below this line, based on the pattern towards unequal negative selection as representated by differential selection, leading to speciation, or "blanket" negative selection which can be considered non-differential or inherently stabilizing selection.

In the graphical representation of this model, we can consider the values applicable to a single normally heterzygously reproductive population, or of two different species that are closely related within a trophic-taxonomic table, such that we derivea normal unmodal bell shaped curve below that illustrates the range of variation found within either a heterozygous population or two closely related populations.

In the model below, we must see that evolutionary development is defined by the limits of minimum and maximum sustainable equilibrium, rather than by the line of optimal equilibrium, such that a population will normally oscillate between these extreme limits within a stable ecosystem. The lower limit line defines the cut-off lines of the curve of normal distribution of a population, beyond which negative selection is expected to occur. The upper limit governs the potential heighth of the curve, and thus indirectly sets the optimal line of equilibrium such that it defines the degree of relative heterogenity or homogeneity (variance or similarity) between two populations or subpopulations.

The four vertical lines represent the limits beyond which selection is expected to occur, two at each peripheral end and two in the center, such that different selectional patterns will result in movement of the lines to the left, right or center.

The center-lines will converge until they come together, or spread apart until they reach the peripheral limits. The total area under the curve represents the total range of trait-variability comprised of either a single intra-specific population or

two closely related inter-specific populations. The upper horizontal line represents the maximum limit of saturation that defines the total capacity of the system. The intermediate upper horizontal line represents the limit of mutual equilibrium that can be achieved by a population or related populations. This line will raise or lower depending on the relative distributions of both overlapping curves. In reality, both lines would be oscillating and would gradually fluctuate, depending on changing external environmental conditions. The lower lines would represent the minimal level of equilibrium or maximal level of disequilibrium beyond which there is a zone of rarefaction. The bottom line represents both the total spatial distribution and the finite limit of a potential ecological vacuum achievable in such a system.

The difference between these models if it were a single specific population or two closely related populations, is that the center zone would be defined in the first instance of a single population as the region of greatest intraspecific competition, while in the second instance of two competing populations, it would be the region of greatest interspecific competition.

If we were to map this distribution over time, we would see a fluctuation of the basic set of values governing these relationships, such that the shapes of the curves, the ranges of overlap and the lines would all change positions. The normal distribution can be considered to by a synchronous or instantaneous cross section of a population that is changing dynamically through time. In any given ecosystem, this would be but one thread in a bundle of similar kinds of threads that are bound like a rope through time.

If we were to return periodically and map our thread, we would yield different profiles of our distributional patterns. If we did this enough, we would find a predictable set of patterns that describe possible pathways of periodic recurrents of such patterns, such that the wave-patterns of the oscillation of the system would be something like the following possible patterns:

Case 1: Stabilizing selection "toward the center" that leads to narrowing the center range.

In the first case of stabilizing selection, increasing competition would lead to selection "toward the center" which would tend to narrow all the cut-offs toward the middle, with the result of elevating both curves up to or even above the limit of total saturation. Equilibrium is maximized in this context.

This state is considered the "start" state in this model because it depicts the total ecosystem in a state of saturated equilibrium, which implies several things, most important of which is the point at which reproductive social selection is held to be of greatest importance (at maximum saturation). In this system, it must be understood that changing values are not automatic and synchronous. Rather, there is always some implicit lag between changes in variable states, such that there occur resonance throughout the system.

In case two, a system that has maximized itself through competitive selection and narrowed its range of variability, is "primed" for destabilization that begins with a directional selection either to the right or two the left. Under such conditions, one "group" or "subgroup" begins winning out over the other, leading to increasing stability of one at the loss of stability of the other.

Case 3: Disruptional Selection, "selection away from the center", results in a "collapse" of the ecosystem, which can be considered to be the state of maximum disequilibrium attained by the system.

In case 3, it is presumed that the loss of stability in either one or the other, with the gain in stability of the other, can lead to a total collapse of the system if the one group is not brought quickly to extinction or displaced out of the system. Disequilibrium sets in due to the imbalance between the two systems, resulting in the loss of stability of both populations. The result is "cladogenesis" or divergence of a single line into two, or else displacement of one population.

Case 4: Balancing Social Selection: emergence of two stable center lines and a movement outward of the extremities

In case 4, the "collapse" of the system will be followed by a reestablishment of balance in relatively rarefied ecological conditions, such that a form of balancing selection favoring two separate central locii develops. The system of selection will favor either establishment of an isocline between two populations, with the possibility of increasing niche-competition between them, or the redevelopment of a single heterozygous population due to displacement of one of the subgroups.

Case 5: Diversifying Selection, "reconvergence of the center," results in a period of maximum diversity within the system as a result of wide tolerance limits to either extreme and a reconvergence to a central region.

Case 6 a: Peripheralizing selection: leading to exclusion or extinction of one group or segment of the population in a state of extreme disequilibrium. This is a case of divergent cladogenesis, or else extinction of both groups, or else phyletic evolution of one group and extinction of the other group.

Final condition 1: A single heterogenous species, extinction of one group or subgroup (phyletic evolution)

In this model, any final outcome is possible at any start condition. Whatever the outcome, there is a return to one or the other original start conditions, or else the cessation of the system.

If we begin with a single heterozygous population in case one, we will end up with either two overlapping cogenitor species (case 3), or the displacement of one species out of the system (case 6a), and the return of the system to stable balance. At the next round, we start back either to start states 1 or 2 depending on the outcome.

This model describes a very basic cycle of life in evolution that is rooted to an ecosystem. It is known that in the fossil record extinction is a common pattern. It is also known that phyletic evolution and cladogenesis are also commonly recurrent patterns.

In this model, diversifying selection is held to be a kind of "inclusive fitness" that leads to maximizing of variability of trait-pattern within or between populations, and can be taken as a period of "niche" radiation within an ecosystem.

Drift and selection against deleterious alleles would exist in every state within the system, describing the more or less random fluctuations of the parametersof the system. In all cases, deleterious alleles would tend to put individual in the extreme tails of the curves, leading to their being selected against regardless of the central patterns of the curve.

Negative selection of deleterious traits does not necessarily result in positive selection of adaptive traits. It can lead instead only to either stabilization or extinction.

Any population may drift about a center at any point, but the effects of drift may be more pronounced at some points, at points of the extremes of maximum equilibrium or disequilibrium, than in the intermediate regions. It is possible to imagine drift setting off a steady-state system into a period of resonance amplifying disequilibrium leading to directional selection favoring one group at the expense of the other.

What is described is a period of rapid change and adjustment, followed by relatively long trends of relative stability, represented by the following:

Exogenous changes can be described in terms of environmental fluctuations that alter the thresholds of the curves, and also as the introduction of a third extraneous species to the system. Introduction of a invader population to a system that describes a single heterogeneous population would switch the system to a two population model. Otherwise, it would extend the complexity of the model to a multi-population system that is implied in a "two-population" start state. Exogenous changes can be introduced at any stage in the development of the system, but it may have different consequences depending on the point at which it affects the system. Introduction of exogenous changes can cause a system in a steady state of relative equilibrium to spiral into disequilibrium.

Evolution as natural selection can be described therefore as an indirect process that is cyclical:

Adaptational competition leads to K states of saturation which leads to reproductive competition leads to r which leads back to adaptational competition.

Adaptational selection leads to inclusive fitness increasing environmental fitness maximizing trait variability and slowing evolutionary rates resulting in K which leads to saturation and reproductive trait selection.

Reproductive selection leads to exclusive fitness decreasing environmental fitness minimizing trait variability and increasing evolutionary rates which results in r which leads to rarefaction and adaptational trait selection.

In order to bring closure to the problem of natural systems theory in biological evolution, two issues remain to be addressed. The first is the suggestion of a set of formulas that might describe the patterns above, in terms that represent a kind of "calculus" of natural selection. The second issue is to describe these oscillatory patterns occuring in ecosystems in reference to other proximate and distant ecosystems that are the source of exogenous changes within the system.

I propose a model of a social mountain-island of ecosystems in a web of life that explains two sets of interrelated patterns:

2. The ability of ecosystems to maintain a "boundary" about itself in relation to coevolving ecosystems that includes the description of this "boundary" as a complex pattern of zonation about "core" regions that structurally represent ecosystemic "centers."

It can be seen that some boundary maintenance mechanism exists to confer stability to any ecosystem, but this boundary is relative and permeable, such that individual species may cross it readily, leading to introduction of exogeneous sources of change into the system.

Several assumptions are made. First, since exogenous change in the total scheme is held to be essentially random from the standpoint of the internal dynamics of an ecosystem, over the long run there should be some relatively constant value of such change, which I will represent as the variable "D". If we consider a point-diversity model, we can see that the value of magnitude assigned to D will be a consequence of the size of the ecosystem. The larger the ecosystem, the greater the perimeter of its boundary and the greater the amount of exogenous change occuring across its perimeter. It presents a bigger "target" for invading species.

It can be concluded that for any stable ecosystem, the amount of exogenous change will be fairly uniform over time, but that the consequence of such change will be a function of the actual relative value of that change and the internal state of the system.

In a system that is on the threshold of collapse, even relatively minor exogeneous change values can trigger a cycle leading into disequilibrium. In a system that is very robust and of increasing stabilization near the level of saturation, even relatively major exogenous changes values can have little consequence in disturbing the system.

Nevertheless, there is hypothesized a kind of "chain reaction" pattern that results in periodic "wave" patterns of exogenous changes that can sweep through a network of ecosystems. Nature in the biosphere may organize itself at even higher levels such that a butterfly effect can be created within such a network. This wave patterns of chain reactions in interconnected ecosystems may account for a certain periodicity occuring in such patterns, and is indirectly tied to the periodicity of the internal mechanism itself.

The island-mountain model can be taken as a relatively bounded area of diversity in a sea of relative homogeneity. It is borrowed from the concept of the island that has been so central to evolutionary theory and is applied as a "mountain" of ecosystemic stability on an epigenetic landscape. It borrows also the concept of the mountain as the terrestrial equivalent of an island that features its own biodiversity of habitat and zonation. In this sense, even a continent can be considered a mountain on the seas. Around the mountain-island can be considered to be an intermediary zone that can be constituted by different possible ecotones. Isoclines would describe zones that range from the intermediary zone to the core region of the mountain-island.

Island models have been important constructs in evolutionary theory and experimentation. Equilibrium theory has been applied to island models.

It is known that larger islands (or island-mountain areas) support more species diversity than smaller ones. There is a linear increase in taxon diversity with increase in island size, in which a ten-fold increase in volumetric area about an island corresponds to a doubling of the number of species in the area. A slope of linear regression through such points is designated as the taxon's z-value in any particular island system. Z values generally range from about 0.23 to 0.33 between different taxa on different isolated islands, and this value becomes the exponent of the following formula:

Rearranging this formula with logarithms, one gets the following linear equation in which z is the slope:

Topographical diversity results in large z values and in spatial replacement of species leading to "islands within islands," while low z values lead to reduced species replacement and relatively homogenous conditions.

It is known that continent "mountain-islands" of comparable size to true isolated islands in general support more species at higher trophic levels than true islands of equal size. The rate of increase of diversity of a continent "mountain-island" increases also with increased area, but the z-value is generally not as great as on a true island, being between 0.12 and 0.17. This difference is held to be due to the relative isolation of islands and the "sampling" characteristics of continental "mountain-islands" where species requiring greater area may occupy the region on a regular but discontinuous basis. Islands cannot therefore support the higher trophic levels found on continents that inherently require greater areas than isolated islands afford.

It has been conjectured that introduction of new species to an island is inversely proportional to the species diversity of an island that is tied to the relative density of the island. The rate of extinction on the island should also increase with the increasing diversity of species on an island.

The invasion of new species is linked in this theory to the extinction of old species. Equilibrium on the island will be reached when the rate of immigration equals the rate of extinction. Rate of immigration (π) and rate of extinction (Þ) largely taked the place of birth (b) and death rates (d) in the previous theories of equilibrium. The number of a species N in the original formulas is replaced by a variable of the species diversity, or S. The resulting equation describes a stable state of dynamic equilibrium.

In the initial development of the formulas, we assume linear variation of the rates of immigration and extinction, such that:

Where π₀ is the rate of change with no species on the island and a and ß represent rates of change of immigration and extinction as S increases. At equilibrium, Sˆ, the rate of extinction and immigration must be equal, such that the two formulae are equal:

This formula is identical to the expression for carrying-capaicty K in the logistical equation:

The average rate of immigration per species (ˉ π) and the average rate of extinction per species (ˉÞ) can be obtained by dividing by the number of species not yet on the island (P - S) and the number already on the island (S), such that:

At equilibrium, rates of immigration equal rates of extinction. This results in continously changing composition of the island (or island-mountains) biotic profile, while the island itself will remain relative stable as an ecosystem. At equilibrium, Sˆ, we get

Equilibrium will increase with increasing P and average rate of immigration, and decrease with the increasing average rate of extinction. The average rate of immigration is the same as the rate of change of immigration (a) and the average rate of extinction is equal to the rate of change of extinction.

Immigration rates are a function of dispersal rates, which decrease exponentially with distance. Rates of extinction are held to be unaffected by relative distance or isolation, but are related to the area. Decreasing areas should result in increasing rates of extinction because smaller areas can support lower levels of saturation and equilibria. Two islands of dissimilar area but equal distance from a source continent should experience different rates of immigration and extinction. Replacement should be more rapid on the smaller of the two areas. Increasing density of areas should also result in higher replacement rates.

Consideration of equilibrium equations tied to immigration/extinction in island models is related to a model of one way genetic flow from a continent to an island that has particular value in considering continental island-mountain models. Gene flow is considered an homogenizing force in evolution that is contrasted to the flow of drift that is the result of relative genetic isolation of two populations. The two forces are held to be counterbalancing and lead to shifting balances or dynamic equilibrium of partially closed systems as we find with all ecosystems.

The effectiveness of gene flow can be measured as the amount of migration (m) and the degree of genetic difference. If pˆ is a frequency constant of a large group and p₀ is a small isolate subpopulation and p₁ represents one generation of gene exchange, then:

The formula determines the amount of gene flow between a continent population and a subgroup population, or vice versa, with each successive generation. The first generation equals:

Subsequent generations can be easily represented in the same formula simply by changing the subscripts, such that the second generation is:

The frequency of the allele of the gene pool does not change. After "n" generations, we would get:

From which we can express the rate of migration (m) as the function of gene frequencies.

Gene flow is expected to be a strong cohesive force in nature that binds populations into single evolutionary units, but gene flow can also act to spread favorable genetic combinations among populations. This leads to a "shifting balance" theory of evolution based on the amount of gene flow, the degree of difference between two populations, and the flow is unidirectional from an almost infinitely large continental population to a small isolated island population.

From the standpoint of island-mountain models, it can be expected that there is a relative "boundary" about an area that circumscribes an ecosystem that defines the relative "distance" betweent that area and other neighboring "source" areas. This boundary is complex and variable, rather than being one of absolute geo-physical distance.

Natural barriers are present on continental systems of course, but the kind of barriers that exist in island-mountain models must be available even in conditions of high diversity and relative homogeneity, as in tropical lowland regions. Such complex boundaries must be defined by the internal dynamics of the ecosystem, relative to the degree of saturation of the system. For any potential invading species, there is some "threshold" of adaptational fitness that it must pass, in order to be successfully enter and adapt to the preestablished system. This I will call the "passing threshold."

This adaptational threshold can be described as the potential counter-adaptational fitness of the preexisting species that are closest to it in resource-profile. If a "niche" is relatively open, or it enters at the periphery of a preestabilished system, then the threshold should be relatively low. If a niche is filled by a stable, preexisting group and the niche defines the center of equilibrium for that group, then the threshold for passing must be relatively high. If a species can pass into a system, over that threshold, then it can successfully adapt, and result in the displacement of species that overlap in its trophic niches, leading either to extinction or displacement of the species.

If entering an island-mountain system requires a threshold value, then exiting the system successful might also require a threhold value, which I will call an "exiting threshold." A group displaced from such a system must either go extinct or leave the system and reenter a new one. Leaving one's place in a preestablished order entails a certain loss of fitness obtained within that order. If a group cannot obtain that "transitional fitness" threshold, that is tied to adaptation in peripheral and transient conditions and reach the "entrance" threshold to another system, then it will be doomed either to extinction, or possibly to a "fugitive" state in some "ecotone" between major ecosystems.

A group that is displaced from a niche in an ecosystem, that must find itself a new niche in another system, is therefore at a fundamental disadvantage in that its adaptational fitness must cross two sets of thresholds that should be considered unconnected and separate. Of course, species invade all the time, with the presupposition that such invasion is a natural reponse to displacement. But it is possible that many invasions are not so much a response to ecosystemic displacement as they are a result of population pressure from a densely saturated system. Any population must regularly throw of its "tail end" population, and some percentage of this must disperse some distance from the core concentration of island-mountain areas to other areas. Either way, the consequence should be the same.

The description of the island-mountain model describes a bounded ecosystem that represents a series of spatially organized habitats that coexist through time within a common area. They exhibit a sufficient level of functional coherence as to preserve the integrity of the system as relatively separate or "isolated" from any other system. To some extent such systems are purely a function of distance. But many other factors may impinge to create a boundary around such a system.

Island mountain models may apply to real islands in the ocean, but they are primarily intended to apply to areas that can be analytically circumscribed on continents. In such a model, I hypothesize that there is one or more "core" areas that define its "gravitational" center. It is the center of balance of the system, and should in theory be an area that comprises a local peak in species diversity and density of saturation. It should also be the region that comprises the greatest heterogeneity of species across trophic-taxonomic categories. In a dense, flat rain forest, a tall stand of trees, or even a single tree, may therefore effectively comprise its own small eco-system.

The core of an island-mountain may in fact be itself so hypertrophied that it can be divided into zones of an inner and outer core.

Around the edge of an island-mountain, and of varying dimensions, should be an intermediary zone of transition that is marked by great diversity of species and possibly by many small peripheral ecotones. This defines the periphery and boundary of the island-mountain, and its intermediate range describes the boundary about an island-mountain.

Between the core and the periphery would probably be one or more isoclinal succession zones that would be marked by increasing degrees of diversity and resource concentration/biotic saturation. Like the periphery that surrounds it, these zones should range from being non-existent to being very vast stretches, and may be variegated in a fashion to create a "patchwork" quilt of sub-systems within the larger framework. This highlights the fundamental relativity of any ecosystem in the biosphere, that it is always a part of some larger ecosystem, as well as a part of the total biosphere.

Continental island-mountains therefore are non-isolated in a fundamental sense that true islands are. They are always a part of some larger and more rarefied ecosystem, or set of ecosystems, in which it plays a part-whole role. Periodic fluctuations of "waves of invasion" can be expected in such ecosystem networks, or biotic webs, which leads to a secondary pattern of "inteference" oscillation in any ecosystem of which it is a part. Such waves can be seen as "evolutionary" surges like "hammers" that sweep through areas, bring disequilibrium and change in their paths.

In such a manner, I have attempted to apply systems theory to an understanding of the basic mechanisms of natural selection that underlie evolutionary process in a consistent way. It is clear that the Theory of Evolution is incomplete. The multiplication of theoretical terms and concepts and the variability of their values in the elaboration of theory are a principle indication that the "synthesis" is yet to be complete. And this is perhaps how it should be. Biology presents an inherent dilemma of being a kind of intermediate theory between the purely physical and the even more chaotic social sciences. It has gained a great advantage in this regard, in having a grand synthesis in the first place. But when we try to nail this synthesis down theoretically to an airtight system of generalization that might be explicable for every instance observed in nature, we get increasing degrees of leakage about the seams.

	K₂/ a₂₁ < K₁	K₂/ a₂₁ > K₁
K₁/ a₁₂ < K₂	Case 3: either group wins	Case 2: group 2 wins
K₁/ a₁₂ < K₂	Case 1: group 1 wins	Case 4: neither group wins

Predator/Prey	Prey increases	Prey decreases
Predator increases	Both species increase	Predator increases/prey decreases
Predator decreases	Predator decreases/prey increases	Both species decrease.

GroupA/GroupB	A (dN₁/dt) > 0	A (dN₁/dt) < 0
B (dN₂/dt) > 0	Both species decrease	A increases/B decreases
B (dN₂/dt) < 0	A decreases/B increases	Both species increase

	Prokaryote	Fungi	Proctoctista	Plantae	Animalia	Total
Geo-Physical	+/-	+/-	+/-	+/-	+/-	+/-
Biomass	+/-	+/-	+/-	+/-	+/-	+/-
Decomposer	+/-	+/-	+/-	+/-	+/-	+/-
Producer	+/-	+/-	+/-	+/-	+/-	+/-
Consumer 1	+/-	+/-	+/-	+/-	+/-	+/-
Consumer 2	+/-	+/-	+/-	+/-	+/-	+/-
Consumer 3	+/-	+/-	+/-	+/-	+/-	+/-
Consumer 4	+/-	+/-	+/-	+/-	+/-	+/-
Total Value	+/-	+/-	+/-	+/-	+/-	+/-