Chapter VI

Stable & Dynamic Biological Systems

Global Models of Co-evolutionary Development

In evolutionary theory, change is explained on several levels. First it is described in gross terms of mechanisms of selection that are assumed to somehow affect speciation, usually applied in the form of population genetics. Then it is also described in micro-evolutionary terms of the actual genetic mechanisms that are involved in genotypic modification of the individual that underlies evolutionary process. Also, in terms of the natural history record, change is described with a broader brush in terms of taxonomic trees and phylogenic speciation. Selection models used in the former cases are usually explained in terms of the observable consequences upon the population, which are differential pathways of speciation, rather than in terms of the causal mechanisms that result in these alternative pathways. Linkages at the macro-evolutionary level that invite eco-systems concepts, and at the intermediate level between micro-evolutionary processes and population dynamics, are not as well defined, nor are the concepts of selection and fitness as these apply on these various levels.

I attempt in these pages to address what I believe to be basic issues in the biological sciences that relate these problem sets to systems theory and to some general synthetic aspects of evolutionary theory. In particular, I've come to focus on what I believe to be basic theoretical implications about our understanding of the related concepts of "fitness" and "selection" that are held to drive the somewhat blind processes of evolutionary development.

I propose these as basically heterodynamic second-order feedback mechanisms that serve to maintain long-term adaptive stability of Mendelian populations, especially in the context of dynamic equilibrium of coevolving ecosystems within a global evolutionary framework.

In this theoretical construction, I propose that many basic models that are applicable to the general descriptive understanding of eco-systems can be applied in an analogical way to co-evolutionary systems. I also propose that, as an essentially historical system with a central theoretical motor, the theory of evolution is fundamentally different from synchronic and functionalistic models found in ecology. The net result is that the outcomes of adaptation in any given eco-system are not necessarily the same long-term outcomes for such a system that is evolving. Since, by definition all natural eco-systems are evolving, it follows that models adapted directly and in unrevised form from eco-systems theory do not fit evolutionary constructions in the same way. In general, they undergo evolutionary transformations that are important to comprehend.

In general, I will make the following basic observations from a natural systems standpoint:

Evolutionary systems are defined by basic geophysical parameters from which they arise.

Evolutionary systems tend towards increasing growth, differentiation and complexity.

Patterns of differentiation and complexity tend to be historically irreversible. (i.e., once one species divides sufficiently into two, the two cannot become one again, in general)

Patterns of growth, increasing differentiation and complexity result in cyclical patterns of periodic alteration and replacement once the basic limits of growth of the overall system have been exceeded.

I propose that there are some basic concepts found in ecology theory that has direct relevance to understanding evolutionary development. Thus we can posit a general equivalence between some concepts in ecological theory and their transformed constructions in evolutionary theory. Underlying this general equivalence is the presumption that the ecological principles that are found applicable in current eco-systems are in general relevant during most if not all periods of natural evolution, though in modified form especially for the earliest stages of the origin of proto-typical life forms.

Some basic concepts apply:

Evolutionary succession: Evolving biotic communities undergo phases of succession in time that are comparable to what is understood in ecosystems. Thus we can postulate pioneer communities, and primary and secondary stages of succession, with possibly intermediate transitions, and the result of a climax stage. We can speculate that this process, which is cyclical, will repeat itself over and over again in different Taxon cycles.

Evolutionary climax: Under stable conditions of optimal adaptive equilibrium, communities will achieve a biotic climax that represents the greatest degree of heterogeneity, differentiation, specialization, complexity and saturation of populations that are possible within the constraints of that system. Such climax communities may be extremely stable systems and relatively long lived.

Evolutionary Regimes: An evolutionary regime describes a stable climactic system in a biotic realm that is dominated by a distinct range of life forms that have a monopoly at the apex of the life pyramid. In such systems, biological constraints tend to outweigh geo-physical constraints, hence the structural patterning of such systems tends to be stable and difficult to alter, though saturation in the long term leads to environmental degradation.

Co-evolutionary systems: Co-evolutionary systems can be described as two or more biotic communities that are overlapping in their adaptive patterns or else contiguous with one another, but are evolving upon separate but interrelated pathways. They evolve interdependently, such that changes affecting one will lead to alterations in the other. Co-evolution is a selection concept of evolutionary ecology that describes the mutual evolution of two interdependent species. Individuals and individual groups may pass between such systems, and occupy places in both, but the directional pathways each system achieves are basically separate. Two systems that are not directly connected may in fact be coevolving systems if resources they share come from a common pool or if some intermediary life-form influences both.

Evolutionary bio-schismogenesis: Borrowing Bateson's famous concept, I will speculate that co-evolutionary systems (and subsystems, i.e., species and groups) will tend towards a pattern of "bio-schismogenesis" under certain conditions.

I will state this model in simple terms that will be reiterated throughout this theoretical construction. For any hypothetical co-evolutionary system, there are at least two subsystems "A" and "B" such that changes affecting subsystem "A" will result in changes affecting subsystem "B" which in turn affect changes in subsystem "A", such that we get a formula of the following general form:

B = X - A(z)

where "X" is some composite value that can be expressed as (zB) and where (z) is some other independent composite value.

Evolutionary Inertia, Acceleration & Momentum: Biotic communities that cohere to constitute stable evolutionary conditions tend in the long run to acquire integrative structural properties that confers "resistance" to change to the overall system. Such systems gain acceleration along an evolutionary pathway and achieve directive selection momentum along that pathway. This often leads in the long run to extinction.

Evolutionary Equilibrium: Evolutionary equilibrium can be defined as a condition of relative dynamic balance achieved within an evolving eco-system and between co-evolving ecosystems such that there is a temporary and extended condition of relative structural stability of relations within the system. Evolutionary systems and subsystems seek this condition of relative homeostatic equilibrium naturally, because as organized energy systems they are working systems.

Evolutionary Entropy: No evolving system or sub-system is perfectly adapted or achieves a state of perfect equilibrium. All systems, because they are changing, and because they are part of a larger system, must deal with a certain amount of evolutionary "noise" or entropy that can be equated with dysfunctional relations, disequilibrium, and instability in the system. Though evolutionary systems always work toward equilibrium, in the larger structure they always tend to decay toward entropy.

Entropy connects any ecosystem with a larger world, and defines that system as one that is fundamentally thermodynamic. In general, noise or entropy will be represented in the system as a measure of "uncertainty" (U) that affects every equation, such that (z) in the equation above is always constituted by some uncertainty factor (U) related to some other set of values (V), such that:

z = V/U

It can be seen that if the value of U increases, the net effect will be to decrease the value of z.

Evolutionary Collapse: All co-evolutionary systems eventually collapse because of changing conditions in other systems beyond their control that result in undermining the bio-geo-physical platform upon which the stability of the system was based.

Because large co-evolutionary systems may come to incorporate multiple coevolving systems on a lower level, there is the possibility that evolutionary collapse can occur spontaneously within the system, from endogenous conditions rather than exogenously.

In such conditions, endogenous systems can act as a trigger mechanism for exogenous changes, and exogenous changes can serve as a trigger mechanism, or catalyst, for endogenous changes that might lead to evolutionary collapse.

I argue furthermore that evolutionary developments of the long run cannot be accounted for outside of a global evolutionary framework, as the presupposition of a global framework sets certain constraints and conditions upon all co-evolutionary communities.

I will state the basic postulates of this theory:

1. At any one time the total biosphere is represented by a number of eco-systems that are composed of one or more biotic communities.

2. All biotic communities occupy one or more eco-systems and are evolving as systems. Biotic communities cohere into evolutionary eco-systems with distinct but relative and transitional boundaries.

3. All evolutionary communities are evolving at differential rates and along different adaptational pathways.

4. All biotic communities undergo evolutionary succession in several stages that result eventually in relatively stable regimes of evolutionary climax.

5. In terms of basic biological and physical constraints, all biotic communities are at least partially open communities. There can be no completely closed eco-system.

6. Because all biotic communities are partially open and evolving, all biotic communities are at least indirectly connected with all other biotic communities, and therefore all are co-evolutionary at least in some minimal way.

7. Co-evolutionary relationships can lead both to adaptational and counter-adaptational selection patterns between different members of coevolving systems that is a function of both entropy and equilibrium.

7. Co-evolutionary relationships tend in the long run to set up patterns of evolutionary development within all communities that result eventually in the anti-climactic destabilization of climax communities and in evolutionary collapse and mass extinction of certain communities, especially at the apex.

8. Evolutionary collapse is rarely complete, and may follow a cyclical pattern of endogenous/exogenous change mechanisms.

9. With evolutionary collapse, room is made within the evolutionary pyramid for replacement of many forms of life from peripheral biotic communities, which leads to a new cycle of evolutionary development.

It is my conclusion that this kind of cycle tends to repeat itself over and over again at multiple levels of analysis, from areas, to larger regions, to inter-regional biomes and continents, and to the entire planet earth.

This bespeaks a basic form of evolutionary relativity of all coevolving systems. This relativity can be interpreted in the following ways.

1. Species all have their own reproductive boundaries. Each species is by definition evolutionarily unique.

2. Species cohere in biotic communities to create distinctive eco-systems that represent co-evolutionary communities that are unique.

3. Each co-evolutionary community has its own structural integrity from the standpoint of its evolutionary pattern as an adaptational system.

4. Communities overlap and cohere into larger systems that develop their own co-evolutionary integrity at multiple levels.

5. The relational complexes that apply to any one species, community or larger system at any one level, do not necessarily apply to other species, communities or systems in a different time or place.

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Speciation cannot only be understood in the context of molecular biology or alternatively in the framework of population genetics. To focus exclusively on the aspects of genetic transmission is to mostly ignore the epiphenomenal and synergistic patterning that life-forms exhibit at all levels of the taxonomic tree of life, as well as how these patterns interact and influence the outcomes of evolutionary development.

To put this issue another way, we can look reductively at any individual organism in nature as a bundle of cells carrying a unique genetic template of design. We can look at a species as a bundle of such templates, attempting to beat the odds in the game of life to achieve some form of adaptive evolutionary success.

But to see the evolutionary game played in only this way is to miss the most important point. Each individual organism is more than the mere sum of its parts, and is certainly something more than just a vehicle of genetic transmission. Each organism represents a unique solution that evolutionary development has arrived at through a convoluted natural history of trial and error to the general problem of survival of life in complex environments.

Thus, it is clear to me as I pet my dog that a year and a half ago this dog was nothing but a fertilized egg cell undergoing rapid cellular differentiation in the womb of its mother. From that miraculous process developed legs, floppy ears, a white-tipped tail, and a somewhat gregarious, if annoying, personality. These things all combined together to form a total entity with its own functional needs for survival, and these needs are quite complex. Evolution arrived at that synergistic pattern in a fundamentally blind game of chance mutation, and it represents much more than just the epiphenomenal unfolding of ontogenetic cellular differentiation derived from the gamete.

In its feral state, that progenitor of my dog had a unique answer to the challenge of survival that life would have presented to it. That proto-dog did not itself construe this challenge in a genetic way, but the genetic outcome was this dog in my lap. It met the challenge by means of behavioral adaptation to changing circumstances, by means of instincts and complex response patterns that were part of its repertory for survival. Now how can the gene know all of that and transmit it?

The solution that evolutionary history has worked out for any organism is as a living system that has adaptive fitness in a larger biotic context that is circumscribed by the boundaries of the kin-group. Furthermore, it has worked out an even grander solution for the behavioral system of the species as a whole.

It must be recognized that when life begets itself anew with each successive generation, the primary challenge of the organism is not reproduction, but survival. If any organism cannot meet the basic challenge of survival, then the issue of reproduction becomes a moot point. The same holds true for the species as a whole.

Thus our first general rule:

A. Life, for all organisms and all species, is about surviving first, and reproducing secondly.

Thus, it is critical to separate the two imperatives in a kind of basic formula for all life that represents the challenge of survival. The first imperative for survival would be represented as an independent variable (I will arbitrarily call it the Biological Imperative Bi). The second imperative is the challenge of successful reproduction and is a dependent variable (what I will refer to as the Evolutionary Imperative, or Ei) such that for any life form there is a hypothetical linear regression relationship:

Ei = Z - (Bi)(xy)

In this formula,

"Bi" would represent any individual or number of related individuals, Variables (xy) would be the genetic totipotential of any individual (and by extension, the entire group) as represented by some cumulative value of "fitness".

"Z" would represent all those factors (uncertainty and selection) that would impinge upon the individual and the group to determine the net outcome

"Ei" would be the variable of the net outcome of relative reproductive success, which can be interpreted in this model, as successful intergenerational transmission of (xy).

While this formula is over simplistic and would require a great deal of refinement if it were to work in any serious way, it does illustrate in a fundamental manner both the complexity and the challenge to understanding the basic processes underlying evolutionary history. It demonstrates clearly that the basic challenge of biological survival of life comes before and underlies the challenge of reproduction.

If we were to elaborate this basic formula, then we would see that (Z) is a complex set of possible formulas that are primarily the subject of this digression. "Bi" itself would represent also a set of factors that would comprise the minimal requirements that an organism would have in order to accomplish survival. Variables (xy) would itself be some composite value that we would have to derive for the genetic fitness of the individual or the group, and would involve the Hardy-Weinberg equations at the group level.

To proceed with our theory, I will state that there is an inherent paradox of this formula on several levels, that leads to a basic conundrum of inherent biological complexity in its self-organizational information patterns.

First, Ei above, if it leads to success, is manifest in the form of a replacement organism (or group) that must have values comparable to Bi(xy) in the formula that created it. In other words, there must be a measure of high positive corelational value between the two successive sets of values. Thus, a basic feedback loop is described, such that

Ei = Bi(xy)''

And we get the derivative formula:

Ei = Bi(xy)'' ~ (Z) - Bi(xy)

Where the sign in the second case refers to an equivalence value of sufficiently high positive correlation.

If Ei leads to failure, then we get another set of values, such that there is either a low corelational value between the result and the initial values, or there is a negative corelational value, and thus:

Ei = Bi(xy)⁰

There are basically two alternative outcomes that would be possible, and this would entail a kind of arbitrary decision having to be made on some level as to where the cut-off in alternative values should lie. To a great extent, statistical decision making of the null hypothesis would have to be employed. We get something like the following conditional syllogism:

If Ei = Bi(xy)⁰

Then, (Z) - Bi(xy) = 0

Else (Z) - Bi(xy) ~ Bi(xy)''

There is another inherent paradox as well in this kind of formula, for it is clear that from an evolutionary standpoint, both the individual and the group of which that individual is a member, and ultimately, the entire species that individual represents, must be somehow taken into account.

Thus the formula must be modified in three sets of related formulas that represent a kind of system such that modification of values in the first primary formula will result in modifications of the subsequent derivative formulas, such that:

1. Ei = [Bi(xy)'' ~ (Z) - Bi(xy)] or [Bi(xy)⁰ = (Z) - Bi(xy) = 0]

2. Ei^{(subgroup x)} = Sum (Ei ^{(n of x)})/N, where "n" is each and every member of subgroup (x) and N is the total number of members of subgroup (x).

3. Ei^{(species w)} = Sum (Ei ^{(xn of w)})/N^x, where xⁿ is each and every subgroup of species (w) and N^x is the total number of subgroups of species (w).

We can thus imagine a three-step process that must be reiterated each time we calculate the evolutionary imperative for a species, and if we were to put this into a computer language, we can recommend a kind of recursion formula that is something like the following:

If 3 = 2(recursion)

And 2 = 1(recursion)

Then 3(recursion) = 3 N^w where N^w equals a given number of generations of species (w)

Now this kind of formulaic thinking is deceitful in two ways, and this is the third paradox.

First, it is deceitful because in its oversimplification of variables it disguises a great deal of complexity inherent to the fundamental variables of the formula, much of which must be unknown and probably essentially unknowable. Hence, there is much that would be arbitrary in its systematic application to real examples.

Secondly, it is deceitful because in its formulaic complexity it disguises a great deal of simplicity inherent to the fundamental relationships embodied by the formula, much of which must be unknown and probably essentially unknowable. Hence, there is much that would be overly reductionist in its systematic application to real examples.

In other words, there is much that can be said to be mathematically spurious about this style of thinking about evolution. The purpose of this chapter is to work out a larger theoretical framework in which to frame these formulas and the paradoxes they represent. Hopefully, in the course of explication of the larger theoretical framework I will be able to offer basic formulas for some of the variables embodied in these formulas, as well as a means to simplify an understanding of some of the implicit relationships in the work. We need to get back on track therefore to our main line of discussion, and where we left off was the statement that in a general sense, the biological imperative of survival of an organism, kin-group or species, underlies and comes before the evolutionary imperative of reproductive success.

But in the larger scheme, if we look at the formulas, it is obvious as well that the issues of survival and reproductive success are interdependent in the structural patterning of the long run, such that if survival does not lead to reproductive success, or else results in reproductive failure, then there will not be trans-generational transmission, and hence survival of the species will cease at the terminus of the life-cycle of the individual--i.e., the natural death that all organisms are prone to.

Thus, our second general rule:

B. Basic issues of biological survival and reproductive success are irreducibly interdependent such that they constitute a natural system that circumscribes a critical and inherently complex range of relationships between the individual and the group of which it is a natural member.

The end game of this negative feedback cycle would be of course extinction, and extinction is well represented in the natural history record of evolution as one of the most frequent pathways taken by most species. The several million species estimated to exist today, are the product of evolution that has witnessed the extinction of many billions of other species. In fact, it is true that all species eventually become extinct, just as all organisms eventually die, and there is more than just analogy to this understanding of death and the inevitable extinction of species. In many respects a species is like a '"superorganic organism" or form of life that is represented by a system of adaptive patterning maintained by multiple related organisms. The extinction of species holds vital clues as to the fundamental mechanisms underlying speciation.

Just as there are some very long lived organisms, like Koi fish and turtles, and the rare case of a human being that lives to 120-odd years, there are some examples of long-lived species that appear to be remarkably unchanged for many millions of years. Why many species appear and disappear in the blink of an evolutionary epochal eye, and why a few appear to resist extinction time immemorial, is another set of phenomena the explanation of which is vital to the understanding of the evolutionary mechanisms governing speciation.

Another grand paradox embodied in this second digression about death and extinction, is that all extant species surviving today are by fact of their survival evolutionary success-stories. They are all ultimately descended from a very long line that goes indirectly back to the first proto-life forms 3 to 4 billion years ago, or at least to the Cambrian explosion. None of these lines have yet experienced extinction, though undoubtedly most of their cousins have, and though many will undoubtedly experience it in the next few decades. Life extant today represents evolutionary solutions that have been worked out by trial and error over countless generations over many millions of years of blind evolutionary exploration. These solutions were not arrived at over night, and could not have been forged in a day, or even in a week, or even in a single lifetime.

The upshot of this is that we are all, all of life, fundamentally related. We are all "cousins" however many generations removed. Thus on the most basic level we all have a fundamental sense of connection to life in its myriad forms, and we all have a fundamental sense of total relatedness in the game of life. The entire biosphere exists as an adaptational niche for all forms of life occurring within its fold. We share the same water in our bodies, and exchange the same energy sources and transpire the same gases. No matter what extinctions and speciation may come and go, life as a whole must continue its imperative of survival. Otherwise, the absolutely final end-game will be the total extinction of all Life. I will call this the "doomsday prospect" and will refer to such a critical terminus event as the possibility of "Global Extinction."

Thus, our third general rule:

C. Life forms a superorganic living system as a whole that has as its imperative long-term survival, and of which all species are representative members and play a part.

I will refer to the challenge of survival of all life, as a coherent and extremely integrated total system comprising all living forms, as the Life imperative. I will suggest that in the final analysis, the Life imperative underlies and informs the biological and evolutionary imperatives of all organisms and species in important and fundamental ways. It also informs our thinking about such systems. Life can afford to sacrifice many of its species and individuals in some grand game of Russian Roulette, but it cannot afford to sacrifice all of it.

The calculus of biological survival and evolutionary reproduction and speciation must be framed in the larger context of the challenge of life, as a total, global system, to continue to survive as such, to evolve and develop.

Thus, I will formulate my fourth general rule:

D. Life, as a global superorganic system, comprising all biological and evolutionary relationships of all species and organisms occurring within its global system, creates the total synergistic environmental context for understanding the dynamics of evolutionary process and the mechanics of speciation.

All events of speciation and processes of evolution must be understood in this global context. If we look at the complete biosphere, we can understand certain things immediately. The basic cycles upon which life depends are inscribed in the geo-physical patterning of the earth's surface, including the carbon cycle, the oxygen cycle, the hydrologic cycle, the nitrogen cycle and the various nutrient and mineral cycles found in the earth and seas. To a great extinct, the processes of life evolution over the past three or four billion years have fundamentally shaped the surface of the earth itself, such that it has implicated itself even in the gradual transformation of erosion and geological stratigraphy. It is possible that the geology of earth could have evolved in a complete different way if biological life as we know it had not implicated itself in the process early on. Examples of Venus and Mars bear this out.

This leads to my fifth and final general rule:

E. Life as a global evolutionary system is intrinsic to the geo-physical system of the whole earth, and is part and parcel of this larger natural system such that alterations of this larger geo-physical system of the earth are primary exogenous factors of basic change in the biological evolutionary system of the biosphere.

To apply these last sets of general rules to our original formula, I will suggest that the global evolutionary system defines a holistic set of factors that are constantly operating in the background of an organism's and group's environment, and that serve critically to influence the outcomes for both the organism's (and groups) survival and reproduction. These are in essence complex "limiting" factors that provide feedback to the basic system. Thus, I will substitute for the variable (Z) in the basic formula given above for a variable I call "the set of limiting factors of the Life-imperative" and I will define this variable as such:

Z = Li = (Sum(Lfⁿ)/n)/U

where "Li" can be described as the total system of the Life-imperative and (Lf) can be described as any limiting factor relevant to a given situation of an organisms, "n" is any number of such factors, and "U" is the residual average "uncertainty" factor arbitrarily associated to each of the basic limiting factors based on a set of decision rules, such that:

U = uⁿ/n

Where "u" is the uncertainty factor assigned to each limiting factor and "n" is the same as the number of such factors. Substituting, we get:

Z = Li = (Sum (Lfⁿ)/n)/ (uⁿ/n)

Before going back down to the basic issues of fitness and selection governing organismic survival, reproduction and speciation, I must emphasize one final point about understanding the dynamics of evolution and the mechanics of speciation. However mockingly formulaic we may want to render our scientific theories of these issues, it must be understood at the outset that evolution displays a natural record that is fundamentally historical in its patterning of complexity. Anyone who has examined and studied and researched historical subjects will understand immediately what is implied by this sense of history.

Historical information patterning demonstrates a tremendous inherent complexity of an epi-phenomenal landscape. Numerous instantiations in countless and countless complex contexts defy simple causal determinations and straight-forward cause-effect relationships. As the father of American anthropology would have said, the natural history of the evolutionary records is cosmographic in its descriptive empiricism. A boulder rolling down the side of a hill, perchance dislodged by lizard crawling within a crevice where the boulder attaches to the hill, rolls down the hill in an underdetermined manner. All that we know is that gravity demands its descent in willy-nilly fashion. Perhaps it will hit a smaller rock below, and its trajectory of descent shift ever so slightly that it impacts against a tree, tipping the tree and dislodging even more rocks. An avalanche ensues that destroys an entire village at the base of a hill, where another instance of a dislodged rock of similar proportions might of found a state of stable rest upon a ledge below.

In terms of the natural history record of evolutionary process, and understanding the mechanical and dynamic aspects of evolutionary speciation, it must be said at the outset that any "rules" or principles derived from an analysis of its patterning must be at best general and descriptively paradigmatic in their application. They are like the Indirect approach in military strategy derived from an analysis of military history. They appear to be robust, to apply to most cases in a general sort of way, and yet they lack the predictive outcomes kind of validity we expect especially from our physical theories. In systems theory, the most we can come to expect are general ratios or formulas of expectability of different kinds of outcomes. The classic example in nature is the prediction of earthquakes. Scientists may expect with a very reasonable chance of success that an earthquake of major magnitude will strike in a given region, but they cannot predict exactly when or where such an event will actually take place. Thus, historical sciences lack the precision and degree of accuracy we come to expect of "pure science."

Historical sciences, like evolutionary theory, are therefore fundamentally epiphenomenal as systems theory. They exhibit patterns that cannot be analytically described in terms of physical causal relations alone. As precise as we have become in our molecular analysis of organic life-forms, we are still no closer to a kind of theoretical understanding of Evolution that has the kind of predictive/descriptive precision found in the physical sciences. To rely upon such organic analysis alone is to miss the basic point of biological systems theory, and to be over-reductionist in our explanatory models. We must yield certainty, precision, accuracy and prediction, for greater explanatory power in our general models.

In this way, going back to the formulas of evolution above, I will state that in general, it is appropriate to substitute some kinds of analytical processes found in the physical sciences for others. The result is that a sense of systematicity is retained in our ability to simplify our understanding of extremely complex phenomena, but at the sacrifice of the kind of "certainty" and hence "control" we would like our sciences to otherwise have.

In this regard, based on previous research in the human sciences, I will suggest the following kinds of substitutions that I will seek to apply to my models of biological and human systems theory. These are the following:

1. Where possible, ordinal values of measurement are substituted by cardinal values of relative measurement and if this is not possible then by "nonparametric" values of comparative measurement or "concrete description" (i.e., one rabbit, two rabbit, three rabbit, four...)

In such descriptive measurement, a direct one-to-one correspondence between the descriptive term and the thing(s) it describes can no longer be deduced, without disguising a great deal of instantive variability and non-categorical values.

In other words, one biological organism is not just like another similar biological organism, in the same way that we might say that one hydrogen proton is just like the next, and both have the same discrete values.

2. Where possible, inferred causal relationships are substituted by "causal correlations" as implied by the first regression formula above, and if this is not possible then by "corelational" values that assume some kind of indirect or hidden set of relationships.

3. Where possible, basic principles and laws found in the physical sciences, leading to mathematically precise and predictive theoretical formulas, that are judged essentially "correct" as finite puzzle-solutions to specific problems, are substituted by general descriptive rules and paradigmatic statements that are held to govern similar situations and hold true for most sets of circumstances. These general rules and paradigmatic statements do not lack uncertainty and are not unexceptionally applied to all similar situations in the exact same way.

4. The kind of rigorous and faultless logic that can be ascribed to cosmological and physical theories and statements, for instance in the statement of Equivalence, Symmetry and Conservation, does not apply in the same way in the historical sciences and is instead substituted by a kind of alternative and historical logic that is related closely to what is called practical logic and rhetoric. Underlying this logic is a looser kind of three-value non-dichotomization, and of necessity a kind of modus tolens rationalization of deriving an antecedent from a consequent. There is also implicit a form of deriving an "is" (or at least a "was" ) from an "ought." Informing this kind of logical rationalization is also a form of universal common sense that is based on a theory of natural sets, compared to classical exclusive categories. Thus, theoretical descriptions lack exactly the same kind of parsimony of explanation expected in the physical sciences. Instead, simplification of explanation rests on working and heuristic values of achieved realism in succinct descriptive (nonmathematical) explanation.

5. Finally, going back to the language of description, replacing implied one-to-one correspondences inferable from solid physical descriptions with a kind of one-to-one correlation expected and common place in historical descriptions, with all the implications of analogy and homology, metaphor and interpretativism, we are left with a basic challenge of comparing apples and oranges. This kind of challenge is obvious in evolutionary history, especially when it comes to alternative taxonomic reconstructions.

Where possible, I have sought to replace direct physical comparison and "identity" with a form of statistically based system of assumable "similarity," based on arbitrary but explicit decision rules, and if not possible, with a kind of metaphorical similitude or "likeness" that includes the possibility of statistically estimated "likelihood."

This leads into a form of statistical description and decision making that I call possibilist statistics, and is beyond the purview of this text to explicate fully. I only broach the issue here with the point of emphasizing the ways that we can approach the problem of description and explanation in the historical sciences with at least one eye to being systematically self-explanatory, reliable and consistent.

What implications does this have for our formulations above? However refined I may make them, they are just that, generalist formulations, and not "mathematical formulas" as are found in the physical sciences. If systematicity is introduced to the elaboration and applications of their variables and relationships, this systematicity comes from the five caveats about historical science listed above, and not from a hidden presumption of these being physical science-type equations.

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The central objective of this work is to get at a working model, derived from basic general rules, that sufficiently describes evolutionary process as a form of natural biological information system governing biological and derivative relationships occurring in life.

From this standpoint, the model comes to rest on the explication of values of limiting factors that are held to bear critically upon any particular situation of an organism, determining that organism's outcome for both survival and reproduction. Before proceeding with this analysis, I want to further refine our understanding of a systemic model of the biosphere.

We need to approach such a model from a global perspective first, before narrowing down its principal variables.

Basically, the biosphere has been divided into its main components:

1. Life, the sum total of all living matter.

2. Biogenic matter, all the organic matter that has been formed by Life.

3. "Biocosnic" matter, all the minerals and chemicals formed through the interaction of life with the inorganic, geo-physical earth.

In any given, delimited region of the earth, there is represented a total "biomass" of that region that is constituted by these three sets of factors. These biotic factors all feedback in some manner to the total geo-physical system of the earth. This biomass connects to all other regions of the earth at least through the principle geo-physical life cycles described earlier (oxygen, water, nitrogen, carbon, and other minerals). The principal energy base of most life (with a few very rare and unusual exceptions) is solar radiation. Biospheric (or Gross) Primary Production are values that determine the total biomass for a given region, based on the continuous rate of conversion of basic cyclic geo-phsyical nutrients through solar energy by primary producers through either photosynthesis or chemo-synthesis.

In an absolute sense, except for solar energy, which is relatively constant over the long term (variable only by weather and seasonal factors like time of year, cloud-cover, etc.) all the elements of this basic system are by definition limited and finite, though in actuality for any given area, most of the basic geo-physical elements are effectively unlimited. These elements can become limiting factors under special conditions--for instance, in long periods of draught the lack of water can critically affect the life-cycles of large regions. In general such limiting factors can play extremely important roles in determining the outcomes of evolution.

Thus, for any given ecosystem in the biosphere, there is a minimal set of basic limiting factors that are always operating in the background. These factors can be expressed as Fg (geophysical factors) and include the following:

Fg = (L , O, W, C, N, T)

Where:

L = total available sunlight for a given period

O = total available oxygen in a given period

W = total available water in a given period

C = total available carbon in a given period

N = total available nitrogen in a given period

T = total available trace minerals and elements in a given period.

It is evident that any one of these variables can be further analyzed into sub-varieties, especially in the case of T. Many complex formulas and systems can be derived from these different systems, and each describe important cycles in the bio-geophysical matrix upon which all life depends. It is not within the scope of this present work to present a full account of all these factors.

For any given biomass within a delimited region, we can determine the total amount of each basic geophysical element constituted by the heterogeneous composition of that mass. All biomass of large scale is by definition heterogeneous in composition. The relative heterogeneity of any biomass can be determined by the percentage values of each species and their net by-products within the ecosystem constituted by the region, over a given period, including the total amounts of leftover biogenic material from earlier life-forms contributed to the region.

We can also determine the total energy amount represented by that biomass. We can determine as well the amount of geo-physical material required to maintain the total Life system of that region, a subset of this total biomass, for any given period of time. Similarly, we can derive estimates for its total potential and actual productivity over a given period of time.

It was an observation at least since Darwin that nature has tended to naturally divide itself up into more or less coherent regional communities. These are referred to as "bio-geographical realms." The distributional patterning found between these units, and within these units, has constituted the foundation for evolutionary explanation from the beginning.

Another set of finite geo-physical limiting factors are the actual biotic "zones" that the region comprises. The earth has been divided into a number of basic biotic zones that define the extensive and intensive limits of biotic productivity of the earth. Many of these zones are variegated and overlapping, and have a great deal to do with elevation and relative longitude/latitude as well as relationships to major geophysical structures such as mountain chains, bodies of water, valleys, river systems, deserts, islands, continents, etc. Rainfall and average annual or seasonal humidity are also very important limiting factors. These limiting factors can be described as mostly "set" or fixed factors, and cannot be easily altered. They are in the large mostly "density independent" factors.

The limits of these zones are represented vertically from about 10,000 feet in altitude to about 10,000 feet below sea level, and include the following from high to low where the underscore line represents sea level:

3. Para-biospheric Zone

2. Alpine Zone

-1.Euphotic Zone (0-200 m.)

-2. Dysphotic Zone (200-1,500 m.)

-3. Abyssal Zone (1,500-6000 m.)

-4. Hadal zone (6,000 +)

This list shows the greatest and most basic division of zones of Life, between terrestrial and aquatic regions. In general, the biomass of the terrestrial zones is mostly constituted by plants and animals. The biomass of the aquatic zones is constituted mainly by prototista and animals.

Most of the biomass of the earth is concentrated in the intermediate zones (1 to -1) and this is the region where the greatest amount of primary production occurs. These intermediate zones are really only represented by about a total of 6 thousand feet of vertical elevation, and the very middle zone that has the greatest productivity is in fact constituted only by about 1-2 thousand feet of elevation. This means that a good proportion of the earth's surface is essentially unavailable to most forms of life, and most forms of life must be confined to the most intermediate and thinest regions of the biosphere of less than one thousand feet vertical elevation.

Zones are also represented horizontally in terms of geographical distributions across the earth's surface, and are known as biomes.

A principal underlying this distribution of geographically defined biotic zones is that an increase of latitude is proportionate (1˚ latitude (100 miles)/100 feet altitude) to an increase in altitude, such that for each 10 degrees increase in latitude, there is an equivalent rise in 1000 feet of elevation. With each increase in 1000 feet elevation, or in 10˚ latitude, there is a corresponding drop in 3.5˚ Fahrenheit in average mean temperature. Latitudinal zones are divided into the following:

7. Polar (no life)/Nival

6. Sub polar/Alpine

5. Boreal/Subalpine

4. Cool Temperate/Montane

3. Warm Temperate/Lower Montane

2. Subtropical/Premontane

1. Tropical/(Medial)

To reiterate, in this distribution most terrestrial biomass of the biosphere, including the greatest biodiversity and heterogeneity of biomass, is confined to the region between 1 and 0. This biomass in total represents only less than 1/4 the earth's surface. More than 3/4's of the earth's surface is represented by marine and lacustrian biomass, and the main biomass of this sub-sea level "euphotic zone" extends generally less than a hundred feet below sea level, with the regions of greatest primary production occurring only in the upper 10 meters of this zone.

The ratio of total terrestrial to aquatic biomass is unknown.

If the terrestrial zones presented above were plotted against two other axis represented mean annual bio-temperature and mean annual humidity, then there is a clear breakdown of naturally occurring biomes on the terrestrial surface of the earth:

Terrestrial Biomes include:

Tundra

Boreal Forests

Deciduous Forests

Temperate Rain Forests

Temperate Coniferous Forests

Mediterranean Woodland & Scrub

Grasslands

Deserts

Savanna

Tropical Rain Forests

Swamps & Estuaries

Aquatic Biomes include:

Marine

Coastal or Neritic zones, containing the greatest variety of life

Littoral zone

a. inter-tidal zones

b. bays

c. estuarine mangrove zone--salt wedge

d. coral reefs

 

Pelagic or Oceanic (blue water zones)

Upwelling zones

Nutrient deserts

Freshwater:

Lotic systems:

Small order (1-3)

mid order (5-6)

Large-order (7-12)

Lentic oligotrophic mesotrophic natural eutrophic/cultural

Littoral zone

Limnetic

Profundal zone

Benthic zone

In consideration of the basic bio-geographical arrangement of the earth, it is important to speculate that there is a basic division between terrestrial and aquatic zones, and the dynamics of evolutionary process and ecosystems are somewhat different between these two zones. It is also worthwhile to consider that at any one particular epoch of evolutionary history, the earth naturally divides itself into broad bio-geophysical realms that are defined by land masses and intervening bodies of water. Within such realms, there are distinctive varieties of taxons that define unique systems.

I will put forward what I consider to be a third zone that is of great importance in evolutionary terms and that is unique from either the fully terrestrial or fully aquatic zones, and this is the "intermediary" zones that comprise mostly the coastal and littoral zones of the earth. This zone constitutes for the most part the intermediate regions of the earth between -100 feet below and 100 feet above sea level along the edges of most waterways. This region would incorporate most river systems as well as the coastal areas around most great lakes.

It is my contention that this zone is distinct from either terrestrial or aquatic systems, because within its zone it includes many patterns and process that share both aquatic and terrestrial features.

Besides this intermediary zone, there are two other kinds of geophysical features that are important as distinct from an evolutionary standpoint. These are islands that are located within aquatic regions, and mountain chains that are in terrestrial regions. Islands represent unique systems evolutionarily speaking, both because of their relative isolation, and because of the dominance of an oceanic climate over the island.

Mountain systems are unique also because they frequently have their own zonation patterns with distinctive vegetation levels, in part due to the atiabatic winds and weather conditions. They also often represent important geophysical barriers to migration that serve as mechanisms of relative isolation. In general, mountain systems can be distinguished between tropical mountains and temperate-zone mountains in terms of their vegetative zones.

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Based on this model, I will divide the total biosphere into three basic bio-zones: the terrestrial, the intermediary, and the aquatic, with the notion that the intermediary zone is a transition zone of overlap between the other two. This is represented in the following diagram:

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All life forms extant today have in a sense been "winners" in the grand lottery of life. They are all equally direct (and indirect) descendants of the very first forms of life that emerged from whatever primordial soup may have existed 3-4 billion years ago. For as many winners as we can count today (and no one knows the total), there have been many, many more losers lost in the embedded layers of earth's natural history. All these losers represent in a sense failed experiments at life. The game continues to be played, day in, day out. With each passing year new losers are being added to the lists of the dead and gone.

There is also a sense that it is highly unlikely, even impossible in fact, that there will ever be such a set of conditions that there will be a resurgence of an age of the Dinosaurs. The adaptation of the dinosaurs was specific to an era, an epoch, of evolutionary development that was unlike any other in its patterning. In this grander sense, there is a form of biological relativity such that even if one could travel in time to earlier epochs one would not do well, and it would probably be unwise to plant life forms from one epoch into another.

Life of any one evolutionary epoch is totally interconnected and bound within the biological framework of that epoch, and can exist in no other framework in an unrevised way. All forms of life arrive together in the same epoch and in the same global system through long and convoluted pathways of evolutionary development.

Life in general is a semi-determined and underdetermined system, but it is not an entirely non-deterministic system. Deterministic variables in any life system are complex and multiple, and often form long inscrutable chains, but they are critical nonetheless in the final outcomes.

To begin this digression, I will speculate on the following considerations. In any one regional area, or any one ecosystem of whatever dimensions, we can expect to find one or more forms of life from each of the major divisions of life. Thus we are likely to find operating in any regional biome of the biosphere, and in any ecosystem of such a biome, representatives of the following type of system as depicted in the functional discrimination table and the diagram below:

In this discrimination table, it can be seen that the primary consumer is Animalia, and the primary producer is Plantae, both of which are followed by Protista that exist in an intermediate category between the two. It can also be seen clearly that functionally Prokaryotes and Fungi are inclusive decomposers. We can outline the possible pathways between the five categories in the diagram below:

It is obvious that arrows connect the five major Kingdoms of life in both directions and create 16 different possible pathways of direct relationship between these categories. There are even more indirect pathways, if we consider all the possible chains of relationship that are possible within such a system. And if we see reiterative cycles, the pathways become virtually infinite.

Possible relationships embraced within this system are many. Plants frequently rely on different animals for the pollination and dispersal of their seeds. Many kinds of bacteria can only exist in the gut of some kinds of animals, and most animals rely on the presence of these bacteria to aid in processes of digestion. Plants indirectly rely on decomposition of both plant and animal detritus by fungi and bacteria to produce nutrients and minerals in the soil for their own nutrition. Some form of scavenging animals rely on bacteria in the decomposition process, and in turn help to facilitate this decomposition for bacteria. Animals rely either directly or indirectly on plants for their principle sources of food.

Now it can be said that some animals eat plants. We know that by functional classification systems, plants, some forms of bacteria and algae are all primary producers. Their production cycles tend to be based on photosynthesis.

Animals and protozoa are consumers, feeding either on primary producers or on other animals or protozoa or both.

We also know that some bacteria and fungi are decomposers, in that they absorb organic molecules reduced from dead organic tissue. Decomposers are important for returning the entire biomass of the system back to its original eutrophic state, returning the basic components and nutrients back to the environmental cycles from which they were drawn.

We get the following kind of reorganization of the diagram above based upon functional relationships.

We get a complex eco-system at a very basic level that can be said to hold true for almost any sub-zone of any biome we designate. In other words, this kind of basic model of an eco-system must be accounted for in any natural life-context we can designate. Granted, isolated, relatively species-specific systems do occur, but these are the rare exceptions to the rule, whether we are talking about forests, deserts, plains, tundra, oceans or even human urban environments.

We can replace each of the very general categories with a very specific example from any biotic context, and we can get a clear idea of the basic system involved.

Most systems include multiple species in subsystems within each main category, and these subsystems also play an important and sometimes critical role to the successful articulation of the entire system.

If any one of these components is removed from the overall system, it is likely that the entire system will be fundamentally weakened to the point of collapse.

Any such system must exist in a state of dynamic equilibrium, such that if the biomass of any one category were altered, it would result in the alteration of the biomass of the other categories as well, reflecting something like the preceding chain.

In such a system, the total biomass capabilities are primarily determined by the primary producers. On land, the main bio-mass determinant is the Plantae Category, mostly from vascular plants. In Aquatic environments, the main bio-mass determinant is the Protoctista Category, including Algae and Protozoa.

These constitute the main biomass determinants of the basic system. All biomass of the other systems would be proportional to the total biomass available from the primary determinant. This means that the biomass limitations of the other systems are inherent.

Inter-category relations can be described as complementary and as at least indirectly eufunctional. Relations between different species occurring within each category tend to be defined in a competitive framework, as each species would represent a proportion of the total but ultimately limited biomass that would be available in that category. This entails that no single species or group of species can outgrow its own categorical biomass limits, without upsetting the equilibrium of the entire system. This is especially true for the middle consumer category, as the decomposers in general are naturally self-limiting systems. They tend to increase rapidly to the total available biomass, and then subsequently to die off as they decompose the available sources. The biomass limits the decomposers is always the total biomass of the entire system.

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In the explication of this model, I propose the preceding paradigm of what I call the evolutionary pyramid and the evolutionary matrix. For any bounded biotic community in any area of place and period, we can state that there exists a hypothetical framework of relationships that can be depicted in the following ways. The first framework to consider from above involves the Kingdom Animalia primarily, and includes a ranking based on internal biomass of the individual, or its overall size.

Within any biotic community, there exists an evolutionary pyramid of a range of possible body sizes for animals, such that there can occur within the same community a number of different kinds of animals of different sizes that occupy separate niches within the eco-system. I will call these the distinct "morpho-trophic" levels of the system.

Such a pyramid is based on the total biomass and productivity of the eco-system, such that the larger the biomass, the broader the base of the pyramid and the higher the apex of the pyramid. Such conditions lead to greater differentiation of the pyramid, especially at the bottom levels.

In general, there is a competitive relationship between primary consumers and secondary consumers, such that the body size of the secondary consumer will be a dependent variable upon the body size of the primary consumer. This will lead to a feedback relationship, not unlike what was described above, such that there is bio-schismogenesis between primary and secondary consumers, resulting in the pyramid increasing in its apex to its limits.

The growth of the pyramid will result in increasing differentiation of all levels below the apex, such that there will tend to be multiple eco-niches at each level that are evolutionarily open.

This basic model is fit within a larger model that I will call the evolutionary matrix of an ecosystem that describes relationships between primary producers, consumers and decomposers.

The number of possible morpho-trophic niches represented in this matrix are much greater than depicted, especially at the lower levels. It can be suggested that most life forms occur in the intermediate ranges and the largest biomass would be represented in these ranges.

We can speculate that for any one ecosystem, there will be a comparable type of matrix that is possible within the limits of evolutionary development, such that the different niches described within this system may be occupied by different species.

This type of matrix applies mostly to terrestrial ecosystems, and including, I believe, terrestrial based aquatic systems like swamps, lotic and lentic systems, as well as most coastal systems incorporating the littoral zones and even coral reefs. I make this assertion because I believe that there is a basic overlap between aquatic and terrestrial systems that is represented in the littoral zones of such systems especially. In fact, I believe these littoral zones between land and water constitute their own separate and unique biomes that are the most intermediate and perhaps the most important as far as understanding the substrate of the evolutionary process.

For true aquatic zones, I will propose a similar but modified matrix:

A similar matrix can be constructed for what I will call the littoral/inter-tidal biome, which I take to be unique and separate from either true aquatic systems or completely terrestrial systems.

All in all I will speculate that in each and every matrix, we may stipulate the existence of a "pyramid of life" comprising all five Kingdoms that hypothetically may exist in such a system, that is represented topographically by the following "pie of life" model. In this "pie of life" model the total biosphere can be represented, or any ecosystem occurring within the biosphere at any level, with the positions and relations of all species within the delimited system.

There is in such a system some potentiality apex that will be represented primarily by the intersections between primary producers and consumers, that represents the greatest organismic biomass that can be supported within such a system. Decomposer systems do not intersect with this apex, but remain concentrated in the peripheries. The periphery of this system represents both the entire earth, with all its resources and constraints of the system, as well as the total biomass and the smallest size that living systems may acquire.

From this model, we may speculate that overlapping subsystems may arise in any number of areas, as the result of the rising complexity of the overall system. The following model would represent in very simplified form the overlapping biotic subsystems at multiple levels.

Life-pies within life-pies, co-evolutionary subsystems.

Representation of the biotic "pie of life"

Each subsystem would have some basic configuration of the pie of life, and would have its own potentiality apex in the overall configuration. The minimal requirements for this type of co-evolutionary model to work is the hypothesis of minimal boundaries occurring between the subsystems at different levels, in order that each subsystem would accomplish its own evolutionary integrity.

Boundaries between subsystems are geo-physical and biological. Such systems would be seen as emergent in the total pie of life, such that there is a sense of inter-systemic competition between evolutionary subsystems for basic resources. If we turn this topographical pie of life on its side, we get a representation of the potential pyramid of life that would reflect the overlay of the morpho-trophic grid on top of the pie of life.

Evolutionary pyramids

We may go one step further in this kind of modeling, and project the entire pie of life into a long continuum. The result would be a cylindrical representation where the internal sub-system cones would represent evolutionary climaxes. At any one time, we could draw a cross-section of this cylinder to reveal the profile of subsystems in their various stages of cyclical development. If we took core samples through the side of the cylinder, we would get examples that resemble our fossil record.

The total biomass of the surface of the cylinder may graudually undulate or expand or shrink under generally changing earth conditions. If we could look within the cylinder along its main axis, we would see even more wild oscillations of subsystems that expand and shrink and then disappear to be replaced by another system.

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I have thus come to formulate a basic model of evolution that I believe generally and faithfully represents the processes on multiple levels of analysis, as biotic communities that cohere into evolving eco-systems that exhibit some minimal threshold of integrity and stability as biological systems. It can be readily seen that if if we want to move beyond proximate historical mechanisms describing selection events in evolution, we are bound to get lost in the long chains of life that stretch around the world and through its natural history. The complexity that life presents to us at every level is astounding and sublime in the old fashioned sense that Darwin probably understood it. Darwin did not need "systems theory" to frame his famous theory, and it has withstood the test of time unlike any other scientific theory. The remaining chapters are thus devoted to the explication of the basic mechanisms that underlie this model, "from the ground up."

Natural Systems

2001