Theory of Evolutionary and Ecological Spaces
Almanac: Evolution:Environmental, Demographic, and Political Risks
DOI: https://doi.org/10.30884/978-5-7057-6399-3_06
Abstract
For the past 30–40 years, there have been no breakthrough ideas in the field of the theory of biological evolution that could lead it out of a deadlock in which it has been for so long. Apparently, in this case some non-traditional approaches are required.
The author of the article proceeds from the commonality of certain basic regularities underlying the so-called forms of matter motion (hereinafter referred to as the FMMs), i.e. physical, geological, biological, and social ones, and has already attempted to make a parallel comparison of the basic patterns of biological, social, and linguistic evolutions (Kondorsky 2014a, 2014b, 2021, 2022). For a better understanding of the problems of biological evolution raised in the article, homologous events and patterns of social evolution will be constantly mentioned in this work. One should emphasize that the author will speak precisely about homologous events, as it is of a particular importance. Thus, one can speak of the homology of phenomena and structures between individual FMMs.
The philosophical approach to comprehending and understanding the processes of biological evolution is hardly innovative (Zakharov A. 2005; Lyubarsky 1996; Pozdnyakov 2017). The researchers in this field also use terminology and theoretical constructs related to the theory of language (Vasilieva 2001). Moreover, linguists themselves drew attention to the similarity of biological and linguistic evolution back in the mid-19th century.
An integral part of a theory that can be recognized as developed is a set of axioms – basic postulates or initial concepts and principles, i.e. provisions that do not require any proof (Ravkin and Efimov 2006). The author did not set out an objective to analyze the literature on the issues of biological evolution. The idea is to present the main points of the proposed concept using relevant examples.
Keywords: ecological space, evolutionary space, social evolution, biological evolution.
Evolutionary Space
Each of the FMMs has two main levels, two spaces – of development (evolution) and existence. Biological evolutionary space has integrity in space and time. Each development has its beginning and its end. The integrity of the evolutionary space takes place in time from beginning to end as a single system. Evolution should be considered as a systemically organized process (Iordansky 2004) (although the author of this article puts a slightly different meaning to this phrase). Accordingly, evolutionary space can be compared to an ‘unrolling scroll’ in the process of ‘writing a text’. The word ‘evolution’ comes from the Latin word evolutio which means ‘unfolding’ or ‘opening’. At this point, ‘the scroll is completely unrolled’. Consequently, the processes within evolutionary space have stopped. Integrity in space implies that all evolutionary processes at each stage of geological history within the interconnected continents represented an integral whole, a single system.
Evolutionary space has its own set of categories. The evolutionary process is associated with the level of Phylum, Class, Order (in other words, taxonomic ranks that can be considered as evolutionary). Family, genus, species (as ecological taxa) belong to ecological space. Evolution and speciation are the processes that occur in accordance with completely different laws, in different spaces, in different planes. Within the framework of evolutionary space there are no concepts of species, population, kinship, selection, adaptation, etc.
An archetype is the most important basic category of evolutionary space (Kondorsky 2017c). Each evolutionary taxon has its own archetype reflecting the type of its organization, for example an archetype of mammalian and reptilian organization, etc. One can speak of archetypes at the level of Phylum, Class, Order.
Here, the concept of a taxon at the above-mentioned levels is fundamentally different from that within the framework of ecological space – Family, Genus, Species. In the latter case, Family and Genus should be considered as a unity of species that have a common ancestor. Within evolutionary space, a taxon objectively exists because it is based on an objectively existing archetype (Zakharov 2005; Rautian 2001).
Usually, the concepts of an archetype and structure plan are seen as synonymous and most often in the latter understanding (Lyubarsky 1996; Pavlinov 2011). In our opinion, the concept of archetype refers to a complex of those innovations that lie at the basis of a new evolutionary taxon and characterize its type of organization. For the first mammals, this was the acquisition of a mammalian-type sound-conducting apparatus, a jaw joint between the dental and squamosal bones, soft lips, upper ethmoturbinal bone, etc. (Tatarinov 1972, 1987a). One can speak of subsequent more or less significant acquisitions, yet that does change the point. Here, it should be noted that innovations of this kind have nothing to do with cladistic analysis and its terminology. Using this method at the level of Order or Class only confuses the situation. Moreover, paleontologists are quite cautious about using such approaches (Vorobyova 1992; Rasnitsyn 2002; Tatarinov 2003).
There is no concept of a specific ancestral species (or group of species) within evolutionary space, nor is there a category of ‘species’ itself. Essentially, no real ancestral forms have been found not only for known Classes (or even such fossils that can be considered the closest relatives) but also for taxa at the level of Order (Gabuniya 1969; Nazarov 1991; Savelyev 2008; Simpson 1948). I. A. Efremov (1935) believed that it is virtually impossible to detect the intermediate forms due to their extremely small numbers. Recently, a ‘textbook’ Archeopteryx was ‘disranked’ from the rank of a bird ancestor (Kurochkin 2006). To a certain extent, the expansion of knowledge about fossil representatives of the ancestral taxon does not clarify the picture of phylogenetic relationships but makes it more and more complex and unclear (Tatarinov 1972).
Here, one can only speak of reconstruction (a reconstructed ancestor) (Vorobyova 1992; Pavlinov 2011) as a hypothetical model (namely a model) of the ancestral form. One can observe a similar situation in the case of language evolution. In this case, most experts believe that the reconstructed proto-language, which is usually called a ‘protoform’, or an ‘archetype’, is a hypothetical language system. All attempts to identify the main languages in terms of origin with certain known peoples have come to nothing.
The peculiarity of evolutionary space suggests the presence of reconstructions only at the level of Phylum (but not yet at the levels of Class and Order) or only Phylum and Class (but not yet Order). Mammals, reptiles, and amphibians evolved from hypothetical ancestral forms that did not yet have a Class-level organization – just like lizards cannot be considered the ancestors of snakes (Iordansky 1994). As for birds, it is most likely that their initial forms already had a reptilian organization. Therefore, Aves must have a subclass rank. Moreover, the morphological differences between the Orders of reptiles and mammals are much more noticeable and significant than those between bird Orders. It is possible that the ancestor of the Tetrapods was a hypothetical form that did not have a fish organization (any of the known Сlasses of fish, including fossils). The fanlike appearance of all main types in the Precambrian and Phanerozoic (Ponomarenko 2004; Shimansky 1987) suggests that they were based on the forms (hypothetical models) that did not even have a Phylum-level organization.
In order to understand the real situation, one should study the processes within the framework of social history. During the Neolithic period, the Eurasian ecumene took place (as a unique system) which included Southern Europe, North Africa, and most of Asia, even China. All main inventions and technologies of that time – the appearance of grain crops, domesticated animals, the invention of the potter's wheel, the development of smelting of bronze and iron – took place in the centre of the ecumene, Western Asia. Then all this spread throughout its territory, right up to the periphery, through mobile groups which were the exact opposite of the traditional static community (Kondorsky 2017a). It should be noted that these groups did not leave any cultural traces.
Thus, one can state that the occurrence of the main evolutionary ‘inventions and technologies’ also took place at the level of small mobile groups that did not have a ‘species’ form and population structure and did not leave any paleontological traces. The population (as a form of biological existence) and the traditional community (as a form of social existence) are involved in homological relationships.
Each archetype has its own potential for further development. After the formation of the basic archetype, it was improved over tens of millions of years. Some of the peculiar characteristics of recent mammal groups have not yet been acquired by typical Mesozoic mammals (Tatarinov 1972). This refers to brain development and locomotion. Placentarity appeared only at the end of the Cretaceous period. The specificity of the elements of the basic archetype provides (or rather, should provide) the possibility of further development. For example, the three-part structure of the sound-conducting mechanism removes the restrictions on different forms of the external ear for mammals (Agadzhanyan 2003). The same applies to the mechanism of lung ventilation using the movement of the chest for reptiles (Iordansky 1994).
Today, most evolutionary paleontologists hold the opinion that basic features of the archetype were formed in parallel bundles (Tatarinov 1972; Vorobyova 1988). A. G. Ponomarenko (2004, 2005) uses the term ‘lawn’ of evolutionary branches that evolve during the formation of a higher taxon. A short digression here: as it was mentioned above (based on the concept of two types of spaces), the concept (category) of a ‘character’ can only be used at the level of ecological space. Moreover, under this concept the researchers understand any feature of the phenotype that distinguishes one object from another (Shatalkin 2002; Yablokov 1982). Therefore, when speaking about an archetype, one should use the terms ‘element’ or ‘structure’. There was a process of angiospermization (Ponomarenko 1998), arthropodization (Idem. 2004), and mammalization (Tatarinov 1972), tetrapodization (Vorobyova 2003), crinoidization (Rozhnov 2005). Here, ‘-zation’ can be considered as a general method of aromorphic taxa evolution (Ponomarenko 2004).
The main logic behind the reasoning is as follows. At the level of a certain ancestral group, there appear individual elements (or groups thereof) of the archetype of the future progressive taxon (in different combinations) within its different phylogenetic lines. Such a process is called mosaic evolution (Vorobyova 2003). And only one line, where all the main elements of the archetype are observed, is the ancestral one. Thus, it is believed that amphibians are originated from Rhipidistia (Vorobyova 1992), mammals – from Theriodontia (Tatarinov 1976), angiosperms – from Bennettitales (Meyen 1986).
In fact, despite the fact that cynodonts had almost all the elements of the mammal archetype (Tatarinov 1972), they had nothing to do with them in terms of origin. There is a hypothesis about the origin of birds from Theropod dinosaurs. There was a mosaic distribution of avian features in different theropod lineages, including feather cover, opisthopubis and pygostyle, during the process of ornithization of theropod dinosaurs. However, according to E. N. Kurochkin (2006), fan-tailed birds evolved independently much earlier, even at the level of some archosauromorphs – it might even have been the ancestral forms of this group. Thus, the relationship between birds themselves and ornithized theropods appears to be more than distant.
It turns out that all groups that are traditionally considered ancestral ‘ran empty’ in the process of parallel evolution. This applies to both Rhipidistia and Theriodontia. The scheme, which was developed by E. N. Kurochkin providing the example of birds, can be used to understand the origin of amphibians, reptiles, mammals, and even angiosperms. That is, groups that have undergone the process of ‘-zation’ and are traditionally considered ancestral in relation to individual classes are not, in fact, ancestral. The formation of a new evolutionary taxon begins much ‘deeper’ and much earlier. The appearance of parallelisms cannot be considered as a phenomenon of phylogenetic advance (Vorobyova and Nazarov 1988). The real picture is the opposite.
Here, a thing of a paramount importance should be taken into account. For example, let us consider the process of mammalization associated with the appearance of eight key elements of the archetype (Tatarinov 1987). Most branches had 1–3 mammalian characters, and a smaller number had 4–7 characters (elements of the archetype). And only one branch, which observes all eight elements of the archetype, in theory gave rise to mammals. As we can see, the distribution of elements of the archetype along certain branches has the character of distribution of random variables. This suggests that this process is based on statistical patterns. The formation of characters of a superior taxon occurs through trial and error (Vorobyova 1992), a kind of an odd-even method.
Here, one should once again turn to the social sphere. The basis of modern society is made up of political and economic systems, the laws of which are objective in nature, and are independent of consciousness. In order to be successful, a politician and a capitalist must ‘feel’ these laws and follow them in their activities. We have something similar in the case of evolution. Those branches that followed the laws of evolutionary space to the greatest extent had the greatest ‘success’.
After the introduction of the basic archetype of classes, a very long period has passed. The same applies to the level of Order. The basic type of organization of snakes developed at the beginning of the Cretaceous period and did not change for 110 million years (Iordansky 1994). During the period of stasis, further development of the basic elements of the archetype occurred. By the beginning of adaptive radiation, the archetype can be considered fully formed. Dinosaurs and mammals appeared at the beginning of the Mesozoic era and coexisted for 150 million years (Krasilov 1986). At the same time, for two thirds of their geological history, mammals remained small rat-like animals (Tatarinov 1972). Reptiles appeared in the Carboniferous period but underwent strong changes in the Triassic one (Nazarov 1991). Bony fish appeared at the end of the Triassic period but remained scarce until the end of the Cretaceous (Tatarinov 1972). Gymnosperms appeared back in the Devonian period.
If one takes a social FMM into consideration, then a similar pattern can be observed. Tribes that can be considered Germanic appeared almost simultaneously with the formation of the Roman state and coexisted with it for almost a thousand years. It turns out that at first a basic archetype is formed that defines the German type of social organization. Then, for a long time, the process of its development occurs without the manifestation of noticeable external activity. And only after the Roman empire had completely degenerated due to its internal reasons, the Germanic tribes entered the historical arena and became full-fledged subjects of historical space.
It is not the laws of ecological space that form an archetype. Therefore, parallelism cannot be considered as the result of a response from the side of a common hereditary basis to similar environmental requirements (Mayr 1968). In response to similar environmental requirements, convergent forms are formed. The appearance of snakes cannot be associated with adaptation to a semi-burrowing way of life which contributed to the elongation of the body and reduction of the limbs (Iordansky 1994, 2004). Legless lizards cannot serve as a model for the process of formation of the snake archetype. It is also impossible to explain the formation of Tetrapod amphibian features by the adaptation of their alleged ancestors to surviving in shallow waters with a lack of oxygen (Vorobyova 1992). The formation of an archetype is not associated with adaptation to specific conditions. It is necessary to clearly distinguish between structures that appeared in accordance with the laws of evolutionary and ecological spaces. In this regard, the example of sea turtles seems to be quite telling. During the transition of turtles to sea habitation, a reduction of the shell as an element of the archetype occurred in the process of evolutionary development of the Order. When the descendants of these turtles began to live in the onshore zone, a secondary shell of bony plates started to form on top of the rudimentary one. In turn, the descendants of these turtles returned to the open sea and their secondary shell was subject to reduction. The modern leatherback turtle has two rudimentary shells. It turns out that if the formation of the primary shell and its reduction occurred in accordance with the laws of evolutionary space, then the formation of the secondary shell and its reduction followed the laws of ecological space respectively.
Many paleontologists express displeasure with the difficulties in separating parallelism and convergence (Vorobyova 1992; Tatarinov 1987). Essentially, these two concepts are considered to lie within the same plane (Vorobyova 1980). Convergence is viewed as an independent acquisition of similar characters by non-related species, and parallelism is that of related species (Iordansky 1990). The concept of two types of spaces completely eliminates this problem. It turns out to be just as in the famous saying of Jesus Christ. Parallelisms are the ‘heritage’ of evolutionary space, and the cases of convergence are those of ecological space. Convergent similarity is associated with the similarity of the way of life within ecological space and concerns only the form. Moreover, convergence is not associated with the formation of fundamentally new taxa.
During the transition to ecological space in the process of adaptive radiation, the archetype maintains its stability. Out of the 300 thousand species of beetles, not a single one has gone beyond the boundaries of the beetle organization (Rasnitsyn 1986). As far as lizards are concerned, stability of the structural plan (archetype) of the jaw apparatus is observed in different feeding methods. Transformations of the jaw apparatus were not associated with changes in the external environment (Iordansky 1990, 2009). In Gastropods, the skeleton may disappear secondarily, yet the archetype does not change (Shimansky 1987). A very typical example of this kind is the appearance of claws within hoofed mammals in South America (Simpson 1983). The fact is, the archetype of the hoofed organization of this group has not disappeared. And the introduction of claws which were used for obtaining food should be considered at the level of ecological space as an example of a particular adaptation.
The same acquisition can be an element of an archetype and a partial adaptation at the level of ecological space. Reduction of limbs in snakes should be considered at the level of evolutionary space, while at the same time that of the lizards should be considered within the framework of ecological space. If viviparity in mammals is the most important element of the archetype of higher mammals, then viviparity in certain species of reptiles (Gabuniya 1969) can be considered as a particular adaptation. The same can be attributed to the introduction of warm-bloodedness in dinosaurs (Kalandadze and Rautian 1993; Krasilov 1985). At the level of ecological space, the introduction of the alveolar structure of the lungs in secondarily aquatic Pipidae should be taken into account (Severtsov 2008). In all cases, adaptations within the framework of ecological space, did not get any further development as opposed to similar elements of the archetype within the framework of evolutionary space.
As far as the role of abiotic (climatic) factors in processes at the level of evolutionary space is concerned, some scientists believe that reptiles evolved as a result of adaptation to arid conditions, the drying of the climate that began in the Triassic period (Severtsov 1990). The cooling at the end of the Mesozoic era led to a similar selection effect on different groups of vertebrates and the introduction of a heat-insulating layer, a constant body temperature, and a four-chamber heart (Vorontsov 2004). Indeed, in the mid-Triassic period, the dry climate zone reached its largest size during the entire Phanerozoic era (Budyko 1981) which negatively affected amphibians (Shimanovsky 1987) and made room for the radiation of reptiles. In turn, the development of amphibians after reaching land occurred in a humid climate (Vorobyova 1977). However, one must not forget that biological evolution is an organic part of the process of biosphere development, a component of which is a climate. It turns out that the interaction of biota and climate within the biosphere has its own internal logic.
The Order and the processes within its development are on the verge of both evolutionary and ecological spaces. Here, we have a kind of evolutionary specialization when in the process of development of an Order the morphological changes that characterize this taxon are usually associated with one main organ – for instance, with the development of the skull in case of snakes (Da Silva et al. 2018). Cloven-hoofed mammals can present another telling example. The main points in the development of this taxon were associated with an increase in size from the initial animal forms which in size were not much larger than mice (Agadzhanyan 2004) to modern horses and rhinoceroses and reduction of the toes on their limbs in most various ways. During the isolation of South America in the Paleogene period, the most bizarre forms appeared on the basis of the hoofed organization, nevertheless accompanied by processes of limb reduction (Simpson 1953).
Under no circumstances can one suggest that the leading factor in morphological changes in solid-hoofed animals during development was an adaptation to existence in open spaces of cereal biomes (Modern... 1967). One should not forget that most of the history of the development of this Order was dominated by leaf-eating forms. It is just that odd-hoofed animals had the ability to become herbivores (Simpson 1953). Very similar processes are also peculiar of the development of the Proboscidean Order when all possible options for the transformation of the dental apparatus manifested themselves (Agadzhanyan 2004).
Within the process of Order development, more than 90 % of the emerging forms are of an inadaptive nature. Inadaptiveness means the impossibility of further development within the framework of evolutionary space (Kovalevsky 1960). ‘Leaving’ from here is accompanied by either extinction or ecological specialization, as happened to non-adaptive rhinoceros and a tapir. The final result of the Order development is the appearance of ‘final forms’, the ones that do not require further evolution. In the case of solid-hoofed animals this is a horse, and an elephant in the case of Proboscideans.
In the process of their development, societies go through three main stages – an archaic, a classical, and an inertial one. These same stages are clearly visible in the development of individual Orders which correspond to three successive stages according to A. P. Rasnitsyn's terminology (2002) – inadaptation, evadaptation, and stasis. If we consider sauropterygians, then archaic notosaurs are characteristic of the Triassic period, classical plesiosaurs for the Jurassic period, and the final forms are elasmosaurs for the Cretaceous period (Gabuniya 1969). During the inertial period, the complete extinction of this Order took place. This kind of development is typical for ichthyosaurs, dinosaurs and other Orders.
An Order in the process of its development represents a unique system. The peculiarity of its integrity lies in the development process itself. A system is an integrated whole, the existence of which is determined by a natural process that connects the parts of this whole (Queiroz and Donoghue 1988). Here, it is also appropriate to recall L. Van Valen's law (1973), according to which in order to stay in place, one needs to run all the time. Development within the Order is accompanied by the occurrence of various kinds of evolutionary branches that have different potential for subsequent adaptive divergence during the transition to the ecological space.
Within the inertial period, when development is brought to a halt, inertial destruction of the system occurs. According to the established concept, the main factor in speciation (evolution) involves changes in the external environment. Still, the fact is that the development process in all FMMs takes place in accordance with its own internal laws. However, during the inertial period, when the developing system gets destroyed, the aforementioned factor can manifest itself in the form of secondary adaptive radiation.
The formation of herbaceous vegetation during the Cretaceous period contributed to the introduction of four-legged dinosaurs that fed on it (Zheri-khin 1978; Ponomarenko 1993, 1998). It should be noted that the transition of dinosaurs to quadrupedalism did not affect their basic archetype. The forms with powerful dental batteries evolved, adapted to the consumption of herbaceous angiosperms (Krasilov 1986, 1997). In turn, the increase in floating vegetation in the late Cretaceous period and the general increase in phytomass within lake ecosystems stimulated the transition of dinosaurs to a semi-aquatic way of life (Idem 1985).
The structural plan should be taken into consideration (in contrast to the archetype) as a set of structural features and relative positions of the main organ systems that ensure the normal functioning of the organism as a whole (Vorobyova 1986; Starobogatov and Levchenko 1993). The structural plan implies a hierarchical system (Lyubarsky 1996) of merons according to the S. V. Meyen's terminology (1978). For instance, if a limb can be considered as a meron of the first Order, then the foot is that of the second Order, and the fingers are those of the third Order. A. I. Shatalkin (2002) provides an example of the state of merons of the forelimbs of Tetrapods in the form of a bird's wing and a mole's digging leg. The wing of a bird can be considered as something perfect, characterizing the organization of a bird class (subclass). If one is speaking of a specific wing of a specific individual of a specific species, then in order to obtain a meron, it will be necessary to isolate the adaptive components that characterize this Species, Genus, Family. If we perform this procedure with the digging leg of a mole then we will obtain a meron of the forelimbs of the insectivorous Order. A meron as well as a structural plan should be considered only at the level of evolutionary space as a Class of organism parts (Meyen and Schrader 1976; Meyen 1978) exclusive of the adaptive component.
Once again, one should note that a structural plan characterizes the entire morphological structure of the organism. Simultaneously, the archetype consists of new structures that appeared during the formation of a given evolutionary taxon. Moreover, the elements of the archetype cannot be called (considered) apomorphies since they do not fall under the concept of a ‘character’ which is absent in the system of categories within evolutionary space.
Cladistic analysis can provide real (objective) results only within the framework of the ecological space and its taxa.
Archetype is a term used by Philo of Alexandria who understood it as a prototype, something opposite to the concept of matter (Lyubarsky 1996). Owen's interpretation of the concept of archetype within Plato's world of ideas that precede real things (Shatalkin 2002). It turns out that if a structural plan is to be considered as a material, specific structure, then the archetype in its primary basis acts as something perfect (Zakharov A. 2005). If one considers a specific organism, then one can decompose it into specific merons. But, it will not be possible to single out a specific archetype.
If we take the constructive aspect (component) of processes within the framework of evolutionary space, it has a universal basis common to all FMMs and beyond. If we subject the history of the introduction and development of the car to a thorough analysis, then we will find our own archetype, structural plan, non-adaptive branches, taxa, adaptive radiation, etc.
The concept of a structural plan is closely related to the category of homology which I perceive as the classical Owen's understanding of homology as equality, correspondence of morphological structures as constructively identical units within a certain constructive whole (Mamkaev 2012) – the structures in particular, not characters. Simply put, at the level of evolutionary space the same elements of the structure plan are homologous, i.e. merons. Homologization is desirable at the level of taxa of the same level – Phylum, Class, Order (Shatalkin 1990). Although the eyes of vertebrates and mollusks have a similar structure (Ibid. 1990), they cannot be homologous. Homology at the level of evolutionary space should be considered as a correspondence between structures (Pavlinov 2011), and not characters. Evolutionary homology cannot be considered in terms of structural similarity inherited from a common ancestor (Modern... 1991; Shatalkin 1988; Bock 1973). There is no concept of a specific ancestor at evolutionary level.
Ecological Space
Both evolutionary and ecological spaces are initially the spheres of operation of certain laws. The basis of an ecological space is the process of speciation. Here are some clarifying points. Everything that concerns the development and universal laws associated with it (that are common to all FMMs) refers to evolutionary space. As mentioned above, ecological space is a space of existence. An important concept here is the resource of existence. To a certain extent, ecological space can be considered as space of resources (Markov 1996; Markov and Naimark 1998; Van Valen 1976). Ecological space (in a specific sense) can include that part of the Earth's mantle (biosphere) where organisms can exist and, most importantly, find resources for existence, such as food. Physical factors play only a limiting role here. And that applies to poikilothermic animals. Given the availability of food resources, mammals and birds can exist in the most extreme conditions, suffice it to recall the polar bear and penguins. In meat warehouses, rats form stable populations at temperatures of minus 10ºC (Schwartz 1969).
In the process of Order development, various types of organizations appear. Within solid-hoofed animals there are different types of reduction of the toes; in Proboscideans, there are different types of transformation of the dental apparatus. These types of intermediate and final organization are ‘sent’ from evolutionary space into the ecological one where adaptive radiation of various levels occurs on their basis and the adaptive component of the Family, Genus, and Species is formed.
The form of existence of a species within ecological space is a population. Each full-fledged species has a population structure. This applies to geographic, ecological, and seasonal populations. Species of this kind have sufficient potential to respond to external changes without having to form new species.
In modern biology, the axiom is the thesis that the formation of new species occurs on the basis of populations of other species either in an allopatric or a sympatric way. We believe that population is a purely conservative system. It is the structure that must respond to changes in the environment in space and time without the formation of new species. We can say that the more rigid the population structure of a species is, the less likely it is that new species will be formed on its basis.
As an example, we can take the gypsy moth, the ecology and pheromone communication of which I once studied in the Middle Volga region and in Moldova. The habitat of this species occupies the entire northern and central Eurasia and North Africa. Although the main food species of this species is pedunculate oak, the gypsy moth can damage more than 300 species. In each region, a silkworm forms its own geographical populations associated with the nature of the local woody vegetation – a birch in the southern Urals, and wild fruit trees in Central Asia.
The ecologically isolated Siberian population represents the object of a great interest. Here, the transition of silkworms to coniferous trees was accompanied by a radical restructuring of the butterflies' metabolism and behavior when laying eggs. At the same time, in Tartary, the attempts to feed early instar caterpillars with pine needles resulted in their death. At the same time, throughout its entire range, the gypsy moth morphologically remained Lymantria dispar which does not quite fit with the main provisions of the neo-Darwinian synthesis. Here, one can speak of species that have a powerful internal (namely internal) adaptive potential allowing them to exist within a wide variety of conditions without the formation of subspecies or new species (Kondorsky 2017b).
Gypsy moth demonstrated certain levels of flexibility in its behaviour at the level of geographical populations. In Tartary, eggs are laid in the butt-long part of the trunk, below the snow cover in winter. A visible preference is given to birch, extremely flexible terminal branches of which contribute to maximum dispersion of early instar caterpillars. In Moldova, during a population outbreak numerous depositions can be found in the tree crowns. In Eastern Siberia, females prefer rocks that overhang forest valleys.
In the social sphere, the already formed social systems could no longer serve as the basis for societies of a different type. Greece and Rome could unite other societies under their umbrella, create an empire, transform these systems into their own kind through the processes of Hellenization or Latinization but nothing more. The outcasts were the heart of the new type of social formations. This is the ‘gang’ of Romulus, and the ‘people of long will’ who made Temujin the Genghis Khan, the same applies to the apostles of Christ, and the Ansars of Muhammad. That is, all those whom L. N. Gumilyov referred to as the passionarians (or movers and shakers).
The basis of the adaptive radiation of the Galapagos finches was laid exactly by the ‘outcasts’. On islands located close to the continents, nothing similar is observed (Lack 1949). The fact is, the populations of different species present there (including birds) are an organic part of the mainland population structure of these species as a whole, which prevents any further formation. Another interesting point is also quite worth mentioning. The Galapagos Islands are the migratory route of American swallow and common rice-bird (Ibid. 1949). But these two species did not become the elements of the local fauna since migrating individuals are part of the species population, and are not the ‘outcasts’.
Migration at the level of individual small groups of ‘outcasts’ should be considered as the main mechanism for the emergence of species. This kind of migration should be distinguished from a simple expansion of the species' range through populations themselves. For instance, within the Holocene warming, the forest zone was revived from refugial patches of woody vegetation (Agadzhanyan et al. 2011).
According to E. I. Khlebosolov (2005), animals themselves can change their behavior when moving to a new ecological niche. In this regard, they can already be considered as new species. Thus, the emergence of a functionally new species occurs very quickly. It was precisely according to this scheme that the process of formation of the species of Galapagos finches (and not only them) took place. Here, adaptive morphology is of a secondary nature. Colubrid snakes, representing almost two-thirds of all modern snakes, are ecologically diverse and represented by numerous terrestrial, arboreal, burrowing and aquatic forms while not being very diverse in their structure (Darlington 1966).
It turns out that particular species (individuals) have, as it were, an organization basis and adaptive superstructure. The former has structural levels of Phylum, Class, Order; a superstructure involves adaptive structures at the level of Family, Genus, Species. The species is related to evolutionary taxa by its basis, and to ecological taxa by its superstructure. Specialization itself can only affect the superstructure. According to B. Goodwin (1982), morphological characters are divided into deep stable ones, determining a membership in a large taxon, and superficial ones – the variables having a lower taxonomic rank.
The formation of adaptive morphology occurs, in our opinion, on the basis of ‘natural’ variability which is approximated by a bell-shaped curve and differs in nature from its own mutation. Individuals outside the population and stabilizing selection may have significant deviations within this kind of variability. It is this that serves as the basis for the driving selection in the process of species formation (their morphological peculiarity). Differences in the beak shape of Galapagos finches have nothing to do with mutational variability (in classical sense).
Adaptive morphology is an organic part of the ecological niche. If one considers the relation of animals to material objects (nest, anthill, territory, food) as property (Zakharov 2005), then property in this case should be considered as a component of the species niche. There is a process of development of (one's own) resource space. In relation to an ecological (species) niche, the concept of environmental factors is not applicable. In the process of niche formation, the elements of the external environment to which adaptation occurs become an organic part of this niche.
As far as the nature of the ecological niche is concerned, in this article I hold to the concept of a one-dimensional hierarchical niche (Khlebosolov 2005; James et al. 1984; Wiens 1989). In accordance with this concept, a behavioural stereotype is preserved in different living conditions. It is the method of obtaining food that is the primary factor determining the divergence among niches. Other authors adhere to the similar opinion (Vorobyova and Salomatina 1982; Lehr 1979). Feeding behaviour determines the development of the main morphological characteristics and behavioural instincts of a given species. Species of the same family have a common strategy of feeding behaviour (Khlebosolov 1996, 2005).
In this case, the main concept would be a ‘way of life’. Thus, an adaptation ‘can be defined as a set of morpho-functional properties of a given organism associated with ensuring a certain way of life in specific environmental conditions (or a certain way of using specific environmental resources)’ (Iordansky 1990: 246).
At the level of ecological space, one can speak of ecological parallelism when talking about adaptations to a certain way of life that ensures the existence of a given species. Legless lizards presented independently within eight families (Iordansky 1994). In the evolution of tailless amphibians, representatives of a number of families transitioned to tree life which was accompanied by the development of suckers on their dactyls (Tatarinov 1987a). Ecological parallelism must be considered at the level of a specific taxon. The parallelism of structures that remove excess salts from the body into the environment at the level of marine and desert vertebrates (Ibid.) is not to be regarded as one. This kind of broad understanding of this phenomenon only confuses matters.
Parallelism, however, is of a somewhat different nature associated with homological series in hereditary variability (Vavilov 1987) and based on mutational variability within the species. In this case, one is speaking of characters that are usually of a neutral nature. Here, the closer the species are to each other genetically, the sharper the similarity between the series of morphological characters is. Genetically close species are characterized by similar and parallel series of hereditary forms (Vavilov 1935). In the first case there are adaptation characters, in the second one there are phenome characters. Considering all three types of parallelism at the same level is not entirely correct (Vorobyova 1992; Tatarinov 1987).
All three types of parallelism are based on autonomous manifestation of some hereditary activity. Consequently, one can speak of three levels of heredity corresponding to three levels of evolution – microevolution (within a species), macroevolution (speciation), and megaevolution (evolutionary space), all of these within the framework of a known genotype. As mentioned before, during the process of ‘-zation’ within the framework of evolutionary space the appearance of individual characters of the higher taxon was of a ‘blank’ nature. This results in a kind of ‘output’ of excess heredity at this level. Something similar occurs in the case of environmental heredity. In the examples provided, the primary activity is the corresponding level of ecological heredity forming a certain tendency which is then picked up by selection. At the intraspecific level associated with population genetics, the main cause of mutations is again the autonomous activity of a given level of heredity.
Thus, one can draw a conclusion that all those factors and mechanisms (within the framework of neo-Darwinian synthesis) which are used by population genetics do not go beyond the boundaries of the species and cannot have a decisive influence even on the process of speciation.
During the transition from evolutionary space into the ecological one, the megaevolutionary level of heredity is preserved and the macroevolutionary level of heredity is activated. One can assume that this kind of transition is irreversible as a consequence of the well-known principle of irreversibility of evolution.
The Systemic Character of Biota
Biota is often considered synonymous with flora and fauna (Rautian and Zherikhin 1997). However, biota is basically not just a collection of species but a system of potential adaptive zones of certain ecological forms at the level of individual species, genera, and families. It should be noted that according to Simpson (1953) the adaptive zone is a theoretical construct. These forms should not be confused with life forms. A life form is formed at the level of a specific organism within a specific community. Moreover, many species in different communities can be represented by different life forms (Yurtsev 1976). Here, the ecological form should be considered at the level of a specimen.
For better understanding of the nature of biota formation and the essence of its structure, let us consider a similar situation of the Great Lakes of East Africa using the example of cichlids, which at one time formed more than 1,000 species in Lake Malawi and about 500 in Lake Victoria. During the formation of lake biotas, there was a process of filling what the author of this article calls trophic guilds: phytophages, zooplanktonophages, molluscophages, invertebrate collectors, burrowing species, prey-chasing predators, ambushers, scavengers, etc. (Lekevicius 2009). Moreover, all this is based on one group, one original organization. If one considers ordinary regional biotas, then such forms can be represented by families often based on several types of organization (Orders). An example is the formation of anthophilous birds in the western (hummingbirds) and eastern (sunbirds) hemispheres (Zherikhin 1978). The penetration of hamsters into areas where insectivorous mammals are completely absent contributed to their secondary transition to a protein-lipid type of nutrition (Vorontsov 2004).
In South America, during the period of continental isolation in the Paleogene period, a shortage of large predatory mammals led to the introduction of land crocodiles and running birds of prey (Kalandadze and Rautian 1993; Rautian and Zherikhin 1997). The change in their way of life had a corresponding effect on the morphology of crocodiles. Their heads, which had a high and narrow skull and forward-facing nostrils, looked more like those of dogs.
The absence of other land birds had a very large impact on the evolution of Darwin's finches allowing them to develop in directions that would otherwise have been unavailable. On continents, finches usually do not develop into warbler-like or woodpecker-like forms since warblers and woodpeckers are already present there (Lack 1949). Considering the Galapagos Islands as a kind of mini-biota, among birds there is a structure of needs for certain ecological forms which appeared on the basis of finches. By the way, birds of other genera and even families may have a characteristic ‘titmouse’ appearance (Chaikovsky 1990). In general, under conditions of isolation any form can theoretically occur on the basis of any organization. There is also another subtle point related to the Galapagos biota. There is no lark-like shape which is usually associated with open spaces. Such habitats occupy a small area here (Lack 1949). It turns out that the adaptive divergence of finches was due to the specifics of full-fledged biomes that make up the structure of the local biota. On the Hawaiian Islands, a more complex structure of the biota resulted in greater abundance, greater diversity, and greater morphological differences in local flower plants. A similar situation can be extrapolated to continental biotas.
One can state that it is the biota that provides for a certain type of organization formed in the process of development of evolutionary taxa, an adaptive zone provided with the necessary resources. It is the biota that minimizes possible competition between species. It is the instability and weakness (poorness of species) of the biota on the islands that explains the noticeable competition between species (Mayr 1968). The process of speciation is under the exclusive control of the biota. The element of biota as a system is not species, genera, families but the processes of their formation and extinction. Biota organizes the external environment, like an ecological niche, with the exception of physical factors to which organisms react as thermodynamic systems.
In the social sphere there is a certain concept of redistribution which is based on the distribution of the prizes of life coming from above. In nomadic communities, the ruler distributes grasslands and migration routes between individual clans so that the nomads have everything they need to feed their livestock, and, on the other hand, for them not to interfere with each other's existence. One can say that biota also has homologous redistribution functions.
Biota can be considered as a unique world (Shimansky 1987) which has a systemic nature (Markov and Korotaev 2008). In historical studies, one usually speaks of the ‘Greek world’, ‘Roman world’, ‘Christian world’ which are based on a certain type of social consciousness. Based on the commonality of basic principles that lie within FFM, one can speak of the Triassic, Jurassic, and Cretaceous biotas as well as higher level biotas such as Paleozoic, Mesozoic, Cenozoic ones as the main stages of development of the biological world. In turn, various kinds of regional world-biotas can take place in space. It would be useful to use the concept of ‘consciousness’ as a kind of ‘genotype’ of one or another biota types. The same applies to the concept (category) of ‘language’. The coexistence of cenophytic and mesophytic communities on the same territory is discrete in nature (Samylina 1974). Mesophytic species are alien to cenophytic species, and vice versa. Consequently, one can speak of Mesozoic, Cenozoic, Paleozoic, Pre-Phanerozoic languages of communication between species within communities (Kondorsky 2014b).
Social space can be considered the most important category of social evolution. Each main stage of human development corresponded to a certain type of social space providing a certain way of existence (Idem 2014a). Similarly, the Paleozoic, Mesozoic, and Cenozoic periods had their own type of ecological space which determined the peculiarity of the existence of species during these periods.
It is possible to use the concept of a ‘civilization’ (Paleozoic, Mesozoic, Cenozoic, etc.). V. I. Dal' considered the concept of ‘civilization’ as the consciousness of rights and responsibilities by a person as a citizen. A citizen species in a biological case acts as a member of a biocenosis within which it can exercise its right to exist, and, on the other hand, to fulfill its ‘civic responsibilities’ by forming and maintaining the structure of the community, participating in ensuring the circulation of substances and energy. Greek civilization is usually associated with democracy, Olympics, temples, theaters, public baths. By the way, in both Greece and Rome baths were the most important place for political communication between citizens. But all these social institutions and structures only ensured that the Greeks fulfilled their civil duties. Likewise, within the framework of the Mesozoic civilization the main thing was not what the dinosaurs looked like but what their size was. The main thing is how they fulfilled their ‘civil duties’.
If one considers biological and social evolution as a single system from beginning to end (as mentioned above), then the stage of social evolution before the archaic revolutions in the middle of the first millennium BC (within the framework of the Egyptian and Mesopotamian civilizations) structurally corresponds to the Paleozoic era, the post-archaic period associated mostly with antiquity in the Greek city-states and the Roman state – a Mesozoic period, in modern times – the Cenozoic one.
The transition from one stage of social development to another occurred in a revolutionary way (Kondorsky 2013, 2015, 2016a). The same can be observed in the transition from one era to another. The main result of revolutions is the elimination of the carriers of the old consciousness, i.e. the elimination of the old elite. By elite I understand the individuals responsible for the preservation and proper functioning of this type of social space. Accordingly, one may speak of the biotic ‘elite’ (dominants) responsible for the formation and maintenance of ecological space. The same thing may be observed during the transition from the Paleozoic era to the Mesozoic one (the famous Permian extinction) and from the Mesozoic era to the Cenozoic one. During these transitions, the dominants of all major communities died out (Krasilov 1984, 1988). At the Permian-Triassic boundary, the dominant block of the terrestrial community died out first (Budyko 1981; Sennikov 2004). Dinosaurs were the dominant group of climax ecosystems of the Cretaceous period (Krasilov 1985), i.e. a kind of elite.
The introduction of world-civilizations in ancient times was always preceded by the migration of ‘unspecialized’ tribes. All civilizations of the post-archaic period – Roman, Greek, Iranian, North Indian, Zhou in China – were founded by shepherd tribes. In turn, the process of formation of world-biotas was preceded by the migration of ‘outcasts’ from ecotones. Subsequently, just as country people in the late Roman empire replaced the degraded and depopulated traditional elite, within biotas offsprings of ecotones replaced dominants who had exhausted their resources (Kalandadze and Rautian 1993). Here, ecotones are understood as spaces not untouched by the processes of phylocenogenesis (Vakhrushev and Rautian 1993).
Functional Space
In addition to both evolutionary and ecological spaces, which are closely interrelated (primarily at the level of Order development) one can speak of a functional space (hereinafter – an FS). An FS is a part of the biosphere. In general, biota cannot be called a part or a component of the biosphere. The connection between biota and the biosphere exists at the level of functional activity of ‘living matter’ and the products of this vital activity. Accordingly, one may speak of a functional niche. Each species has an ecological niche within the biota but not each of them has a functional niche. The ecosystem acts as a functional system. An element of an ecosystem is the functional activity of organisms, and not the organisms themselves. One elk in the forest-steppe is a functional element of several small terrains as ecosystems. Of course, one cannot speak of any population as an element (component) of an ecosystem (as it is often mentioned in ecology textbooks).
Leaf-eating insects can form functional niche during irruptions (population outbreaks). Mass reproduction of leaf-eating insects stimulates the processes of mineralization of leaf and branch shedding and contributes to a more intense flow of the biological cycle as a result of the rapid release of a significant amount of matter and energy contained in the forest litter (Zlotin and Khodashova 1974). At the same time, during periods of depression between population outbreaks, the density of gypsy moths can decrease by 2–4.
It is essential to distinguish between the concepts of organism and specimen (Kondorsky 2017b). Among invertebrates, there is usually a separation of these biological systems. In Lepidoptera, the caterpillar can be considered as an organism in its pure form, and the butterfly as a specimen. In Vertebrates, there are two systems within one specimen, two components – a specimen and an organism. An organism is a thermodynamic system where functions produce the required result with the least energy consumption (Grodnitsky 2002). Different flight modes of birds are characterized by speeds at which total energy expenditure is minimal (Hedenström and Alerstam 1995). The same applies to walking (Minetti and Alexander 1997). At the same time, an organism is a biochemical and biomechanical system.
Adaptations at the level of the organism as a thermodynamic system are situational in nature associated with the action of physical environmental factors. The complete dominance of the specimen as a component of the homeothermic animals forced them to spend more energy to maintain body temperature in unfavorable conditions at the level of organism compared to poikilothermic animals (Iordansky 1994).
The same specimen in vertebrates is an element of a species population, and as an organism, it is an element of a community. Let us go back to the gypsy moth. In the event of a population outbreak, one may usually speak of population dynamics. However, this is not entirely correct. An element of a population is a specimen. Here, the change in numbers affects caterpillars, i.e. organisms. It would be more correct to talk about the dynamics of the gypsy moth caterpillar community. During an outbreak, changes concern not only numbers but also qualitative indicators characterized by the organism – fertility (more than two times), sex-ratio factor, even the nature of pheromone communication. During the period of depression, these parameters returned to their original state.
Marine invertebrates (who have completed the process of evolution), with the exception of arthropods, have already completely transitioned into the functional space at the beginning of the Mesozoic era, where the process of speciation was determined by the needs of the geochemical cycle, and not by the needs of the biota within the framework of ecological space. Even in the Vendian period, the process of colonization of the ocean bottom over vast areas was underway. The rise of nutrients from depth of the ocean ultimately contributed to an increase in the energy available to heterotrophs (Fedonkin 1987). In this case, the biota is formed and exists according to the laws of functional space rather than the ecological one. Once again, it is crucial to remind that the biota within ecological space provides adaptive radiation of all branches in the process of evolutionary taxon development. There is a process of adaptation that ensures the existence of a certain species. Thus, the process takes place from the ‘point of view’ of the species, not the ecosystem. Within functional space, everything happens the other way around. Both evolutionary and ecological spaces are interconnected and form a unique unity of the biological FMM itself, in contrast to functional space, which is part of the biosphere. Speciation is possible, just as the nature and structure of species within ecological and functional spaces must differ. The biota within functional space is close to the Gaia hypothesis developed by J. Lavelock (1979).
The close connection between the processes of speciation and extinction and the nature of the geochemical cycle within functional space is clearly visible in the example of restructuring of marine ecosystems. During the Cretaceous period, as a result of the invasion of pioneer species of planktonic organisms, the rate of pelagic sedimentation increased. Increased accumulation of organic matter in sediments has changed the structure of benthic communities, thus stimulating the development of organisms capable of using these food resources. There was a shift towards mud eaters which had a negative impact on the communities of filter-feeding organisms and recliners, i.e. unattached and immobile bivalves. The relationship between epifauna and infauna changed which resulted in the introduction of forms with a real exoskeleton among the epifauna (Krasilov 1985). The exceptional role of invertebrate organisms within functional space is indicated by the fact that filter-feeding organisms filter the entire water surface in the world's oceans in just twenty days! (Lapo 1983).
In the 1970–1890s, the scientists of the paleontological institute in the USSR Academy of Sciences have done a lot of research on the crises of the Mesozoic biota (Zherikhin 1984, 1987; Kalandadze and Rautian 1993; Rautian and Zherikhin 1997; etc.). Particular attention was paid to the mid-Cretaceous crisis, the largest in the entire Meso-Cenozoic history (Rautian 1997). According to V. V. Zherikhin (1978, 1987), the basis of this crisis was the process of displacement of mesophytic communities by angiosperms. Another crisis also occurred in the Jurassic period (Kalandadze and Rautian 1993; Barskov et al. 1996). The crises at the turn of the Paleozoic, Mesozoic and Cenozoic eras have been studied quite thoroughly.
Here, there is an important point to note here. The latest crises are somewhat different in nature from the Cretaceous and Jurassic ones. In medicine, a crisis is understood as a turning point in the course of a disease leading to a sharp improvement or deterioration of the patient's condition. Thus, the crisis can be considered as an external manifestation of regulatory processes. During the Cretaceous and Jurassic crises, there was no destruction of biota, unlike that on the verge of the Mesozoic and Cenozoic eras.
A number of authors believe that biotic crises are associated with the destabilizing effects of climatic factors, volcanic and tectonomagmatic activities (Krasilov 1981, 1987; Alekseev 1989). At the same time, others associate them with internal biotic factors (Zherikhin 1978; Kalandadze and Rautian 1993).
Here, it is yet again appropriate to turn back to the processes within the framework of a social FMM. In Roman history, the era of civil wars of the 1st century BC shook the foundations of the state. However, this crisis contributed to the formation of a new system of empire governing which ensured its existence for another 400 years. The main cause of the crisis was the degradation of the polis and the polis management system in the conditions of the vast territory that the Roman state occupied at that time.
Something similar happened in the Mesozoic era. It can be assumed that, by analogy with a social FMM, that in the first half of the Mesozoic era the structural basis of the biota was associations of polis-type communities that occupied a noticeably vast territory. These structures had their own internal life and were distinguished by their ecological-cenotic isolation (Kurkin 1976) in relation to other policies. During this period, ecotones occupied (most likely) about half the territory. Paleontologists note the predominantly semi-aquatic nature of communities in the Mesozoic period (Krasilov 1997). The main type of food for dinosaurs was the plant mass of mainly floating plant-bacterial mats (Ponomarenko 1998). This determined the local nature of their habitat (Popov 2005).
In the Paleozoic era, this trend was even more defined. In the Carboniferous period, the vegetation cover occupied a small part of landmass. These were mainly forests of marshy areas adjacent to the seas (Davitashvili 1977).
As remarkable as it is, during the period of social evolution corresponding to the Paleozoic era, the civilizations of that time were located in the swampy floodplains of rivers. The Mesopotamian one was located on Tigris and Euphrates, and the Egyptian one – on Nile. The main structural unit was the nome, which could occupy a noticeable territory but was perceived as a single household, like a house-estate. During this period, there was no concept of a state. All of Egypt was considered a single House. By analogy, one can assume that in the Paleozoic era, plant communities that could occupy significant areas presented a single ecosystem.
In subsequent periods of the Mesozoic era, types of ‘polises’ appeared, distinguished by external activity and ‘passionary’ forms that were open for migration from other, more conservative polises. Associations of these structures are formed in the form of peculiar ‘empires’. Sooner or later, degradation of the polis basis of the Mesozoic biota begins which leads to a crisis. Already within the period of Roman empire, only a shell remained of the policies. In the Cenozoic era, the basis of the structure of the biota became a continuum of communities (Mirkin 1985, 1990) in the form of the biomes that are currently known to us. For instance, the majority of typical taiga plants in Eurasia have a continuous distribution throughout the entire length between the shores of the Atlantic and Pacific oceans (Tolmachev 1986). Cenozoic biomes are formed on the basis of internal colonization, i.e. the maximum development of all land areas. For this purpose, angiosperms represented an ideal tool, especially in terms of their anti-erosion capabilities (Ponomarenko 1993). A similar process of internal colonization in Europe at the beginning of the 2nd millennium AD predetermined the advent of the New Age.
A community is usually understood as a historically established complex of species (Dlussky 1981). P. Giller (1988) considers a community as a group of cohabiting populations. Let us consider the situation using the example of a community of an early spring oak complex which I once encountered in northern Moldova (Kondorsky 2017b). The complex consisted of more than 30 species of leaf roller moths, geometrid moths and cutworms. Thus, here one can witness a species biocenosis, the structure of which could vary from year to year and within specific terrains, but on the whole remained stable. In contrast to the concept of biocenosis as a hierarchical structure (system) of certain species in a certain region, a collection of organisms (precisely organisms, not specimen) within one tree, where contact between them is possible, is exactly what a community is. The community consists of separate specific organisms. It is between these organisms that competition can exist, and not among the species that are part of the biocenosis. The elementary biocenosis here is a terrain, within the boundaries of which there are populations (Gilyarov 1954) of species included in the early spring complex.
Most complex types were content with a ‘small share’ in terms of resources. One can observe a similar situation in the tropical forest where the overwhelming number of species as shade-tolerant forms can be content with the lack of sunlight (Hubbell 2005).
I have established a pattern that if the emergence of caterpillars of the early spring complex coincided with the bud stage of oak leaves, gypsy moth caterpillars, which began feeding later, could find themselves lacking a food resource. This led to the suppression of outbreaks of gypsy moths in certain areas. However, this does not mean that there was competition between species of the early spring complex and the gypsy moth. One cannot speak of any displacement of the latter. Once again, it is important to repeat that competition does not exist between species but between organisms of species (not specimen).
There are many theories of extinction due to climate change and cosmic factors. A detailed review of the literature on this issue is available in V. I. Nazarov's (1991) and L. Sh. Davitashvili's works (1969). In the West, popular hypotheses are related to the catastrophic impact of cosmic factors and climate change (Catastrophes... 1986; Cherfas 1984; Raup and Sepkoski 1984). However, one should note that the factors of one FFM cannot be the cause of phenomena and processes within the framework of another FMM. It is necessary to consider the processes of extinction at the level of ecological and functional spaces and in accordance with the laws of these spaces. As is known, any development processes within the framework of all FMMs have a beginning and an end. It is the process of development of evolutionary taxa (primarily at the level of Order organization) that presupposes the processes of extinction of various kinds of lateral branches.
The groups of organisms that have completely transitioned into functional space can no longer exist in the form of objective taxa of evolutionary and ecological spaces. Any classification here no longer has an objective basis but a subjective one. There are no processes of development of evolutionary taxa and formation of ecological taxa in the process of adaptive radiation. At the level of a functional space, some groups are replaced by others in accordance with the needs of the geochemical cycle. In the Triassic period, gastropods were replaced by the groups of slugs, and by ammonites and nautilids at the end of the Cretaceous period (Tatarinov 1987).
The Factor of Geographical Space in Biological Evolution
In biological evolution, the factor of geographical space is of great importance. The evolution of higher taxa can only occur on the maximum territory characteristic of a given geological period. Even S. S. Schwartz (1980) drew attention to the fact that large-scale evolution and formation of older taxa occurs on continents, and not on islands. In the Tertiary period, the formation of the main placental Orders took place on the territory of the northern continents (Asia, Europe, North America) forming a single whole in the first half of this period (Agadzhanyan et al. 2011). A similar pattern is characteristic of taxa of Class and Phylum rank in previous geological periods. In the Paleozoic and Mesozoic periods, biotas located on the periphery were weak generators of fundamentally new plant characteristics and, accordingly, new high-ranking taxa (Modern... 1988). Angiosperms appeared in Central Asia (Zherikhin 1978, 1987; Ponomarenko 1998) in areas with noticeable seasonal fluctuations (Stebbins 1974).
At the same time, in South America, which was isolated in the Paleogene era, evolution was of a completely different, one might even say ‘horizontal’, nature. Here, on the basis of the ungulate (solid-hooded) organization (archetype) there developed the forms similar to rodents, camels, and elephants, but not related to them (Darlington 1966). Evolutionary taxa can develop normally only on the maximum territory for a given geological period.
The isolated position of the New World similarly affected the languages of the Indians of South America which are distinguished by their extraordinary genetic fragmentation and account for more than a hundred families and isolated languages. For reference, only seven main language families cover 90 % of the population in Eurasia. Social and historical development in the New World, in contrast to Eurasia, was characterized by the absence of the wheel, potter's wheel, horse-drawn transport, bronze and iron tools.
Asia might be considered the centre of origin of the main placental groups. North America and Europe in relation to Asia were biographical dead ends (Agadzhanyan et al. 2011). The introduction of mammals from Asia was the main way of modernizing the North American and European biotas in the Early Cenozoic period. Despite the fact that the connection between Asia and North America was carried out through Bering land bridge which was located on the periphery of the continent and regularly disappeared, all main groups that occurred in Asia were represented in America. They presented something like ‘interconnected vessels’. At the same time, the migrations of new groups were more of a spreading nature.
In Eurasia, the exchange of faunas occurred through ‘ecological corridors’– open landscapes from Eastern Mediterranean Europe through Asia Minor, the Middle East and Northern Arabia to Afghanistan and China (Ibid.). The most interesting thing is that the process of dissemination of the main technological innovations that appeared in the Neolithic period in Western Asia (the centre of the Eurasian ecumene) followed exactly the same path. They spread from the centre to the periphery of the ecumene, particularly China (Kondorsky 2016b). In each region, the forms they had taken had certain local peculiarities.
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