Chapter IX

Evolutionary Dynamics of Ecosystems

Environmental Fitness & Adaptive Selection

by Hugh M. Lewis

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 sub-groupings. 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 behavioral 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 "counter-adaptational selection," as the selection process impinging on other organisms that are the result of the adaptive responses of an individual organism to its own selection constraints. These as a net consequence rebound upon the organism's own selection patterning in a dynamic feedback process of mutual adaptational constraints.

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

1. Evolutionary systems are energy-based systems that reproduce themselves.

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.

2. Evolutionary systems have entropy..

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 succession 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.

5. Evolutionary systems always differentiate and never amalgamate.

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, in spite 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, co-evolutionary 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 sub-evolutionary systems within its boundaries.

All co-evolutionary systems together make up the biosphere.

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

Just like the co-evolutionary 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 cost 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 multiplies 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, ambi-sexual and bisexual in its patterning. The ambi-sexual variety has sprouted little "roots" that allow it to remain attached to the sides of the tide-pools and survive conditions of drying and re-submersion. 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 ambi-sexual and bisexual varieties have come to invade the inner ring of tide pools, and the ambi-sexual 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 asexual 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 mountain 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 happened 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.

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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 super-saturation it is expected that the density-dependent and therefore stabilizing nature of these relationships no longer hold, at least in a relativistic sense. In such frameworks, things socially fall apart.

Understanding this process entails the following presuppositions:

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 evolutionary 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 co-evolutionary 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 inter-specific, 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 competition. 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 clear-cut 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 "counter-adaptational" and co-evolutionary. 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 inter-specifically oriented.

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A one-to-one correspondence between a gene and a trait is a naive and over simplistic 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 poly-morphic or pleio-trophic, 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. This might have two kinds of consequences in the outcome.

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 genotypic 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 genotypic and phenotypic 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.

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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 organismic functioning of an individual of a species

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

Super-systems models, involving regional and interregional adaptive regimes

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 contextually 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.

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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 reproduction 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 super-system 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 macroscopic 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 transferable 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 wild lands 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.

From this standpoint, we can consider the following:

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

Mass extinction as the natural outcome of super-systems and evolutionary aegis or regimes.

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

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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 Lamarckianism. 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 population 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 organismic tree structure is related to all other organismic tree structures on some level or another.

The larger level of evolutionary 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 organismic 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 organismic level, to a population--inter-specific level, up to a global level.

In this regard, informational patterning of Life, and the inferable 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 organismic 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 organismic 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 extra-cellular 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--homeo-thermic 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 sulphrous 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 environments 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 intra-specific 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 predatory ones, for instance, snakes, actually spread out into wide areas as individuals, and only re-aggregate 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.

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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 mutualist at the same time. To the extent that eco-systems are closed affairs with finite energy 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 mutualist 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 complexly organized. These tend to emphasize the survival and reproduction 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 eco-systemic-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 counter-adaptive bio-schismogenesis between interacting groups or populations. The natural outcome is for super-saturation 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 interaction correlates of "r" and "K" in basic population and speciation 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 occurring 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 overpopulation, 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 cris-cross multiple eco-systemic 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 occurring 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 eco-systemic 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 eco-systemic 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 eco-systemic 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 speciation 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-off 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 selection 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.

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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. Counter-adaptation and co-evolutionary 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 heliotropism 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 eco-systemic 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 contributor 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 population 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-occurring is given by a pair of differential logistic equations:

D N₁/dt = r₁ N₁/((K₁ - N₁ - a₁₂ N₂)/K₁)

D N₂/dt = r₂ N₂/((K₂ - N₂ - a₂₁ N₁)/K₂)

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 inter-group competition, the competition coefficient and the N factor of the other group in each equation equals zero, both populations would grow sigmoid 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:

(K₁ - N₁ - a₁₂ N₂)/K₂ = 0 and N₁ = K₁ - a₁₂ N₂

(K₂ - N₂ - a₂₁ N₁)/K₂ = 0 and N₂ = K₂ - a₂₁ N₁

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, Eric; Evolutionary Ecology, 1978: 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, neither 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:

D N₁/dt = r₁ N₁ - r₁ N₁ - ß₁₂ N₁N₂

D N₂/dt = r₂ N₂ - r₂N₂ - ß ₂₁ N₂N₁

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 intra-specific self-dampening rate and the third term the inter-specific competitive inhibition of the rate of increase.

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

D N_i/dt = r_i N_i { K_i - N₁ - (∑ⁿ_/(j≠i) a_ij N_j)/ K_i }

Where i and j subscript species ranging from 1 to n.

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

N_iei = K_i - ∑ⁿ_/(j≠i) a_ij N_j

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.

The predation equations are:

D N₁/dt = r₁ N₁ - p₁ N₁ N₂

D N₂/dt = p₂ N₁N₂ - d₂N₂

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 N₁ N₂. 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 /p₁ and 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:

D N₁/dt = r₁ N₁ - (zN² ) - ß₁₂ N₁ N₂

D N₂/dt = ý₂ N₁N₂ - ß₂N₂²/N₁

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 occurring.

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 inefficiency. 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 optimum 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 approach 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 counter-adaptations 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 super-saurs reached the zenith of their K-development. In such a hyper-developed system, this would be an expected outcome.

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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:

D N₁/dt = r₁ N₁/((X₁ - N₁ + a₁₂ N₂)/X₁)

D N₂/dt = r₂ N₂/((X₂ - N₂ + a₂₁ N₁)/X₂)

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.

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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 constantly 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 intra-specific competition by individuals within one group, which often leads to opposite consequences. In general, intra-specific competition is held to affect the population's tolerance, its increased use of broad-based resources and wider phenotypic variability. Whereas inter-specific competition is held to generally restrict a group's range of resources and habitats, intra-specfic 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 intra-specific 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 intra-specific 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 intra-specific competition may be adapted directly from formulas for inter-specific competition, if we assume that populations can be subdivided along lines of clinal variation represented by their fitness 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 intra-specific competition will lead evolutionarily to selection patterns that result in congenic inter-specific competition between closely related species.

Patterning of intra-specific 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 super-saturation, it becomes a "war of all against all."

It is held that intra-specific 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, intra-specific competition has different selection effects on groups than does inter-specific competition between groups, the latter being restrictive and inhibitive, the former being diversifying and expansive.

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

In fact, it appears to be the case that conditions of high inter-specific competition can result in patterns of relatively low intra-specific 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 speciation patterning that reflects diversifying and balancing selection, and reflects increasing intra-specific 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.

Natural Systems

2001