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G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I http://www.icfcst.kiev.ua/VASECHKO/Vasechko.html [email protected] 16_Axiom6-1.doc AXIOM 6(1) SETS of EFFECTIVE ENVIRONMENTS of PLANT PESTS In this section, it will be given the composition of EEs for groups of herbivores with similar life strategies at combined effects of the levels of ESPPs, the categories of biomass of dominants, and W.C. Cook’s zones of the species range that determine the value of density, patterns of density-dependence, longevity, potency and evaluation of the effects in ecosystems of diverse types. The matter is brought together in the Tables 36 and 37. It should note that the below suggestions as to composition of EEs for the groups of herbivores with close life strategy are extrapolations of data for separate species, as it was done in the Section 5(1). Further, EEs of some groups are concidered now in the first time on the base of very limited information. Therefore, the compositions are less correct than those given in the Section 5(1). Thus, this discource is an initial attempt of generalization of the sets of EEs on the base of life strategies of herbivores. 6.1.1. BIOCENOSES 6.2.1. The level of ESPPs 3.1. "Proper control" 6.3.1. The category of biomass D - I An operation of Antibiosis (and Nonpreference in stem borers) to herbivores. Conductive tissues of stems, branches of trees and shrubs, shoots of coniferous species, matured roots, needles in evergreen coniferous species, and buds. Stem borers, defoliators of evergreen coniferous species and other herbivores, which pretend on host-plant tissues with developed Antibiosis. 6.4.1. Life strategy: Stem borers The group of stem borers includes numerous species of the orders Coleoptera (families: bark beetles, Scolytidae, long-horned beetles, Cerambycidae, buprestid beetles, Buprestidae, weevils, Curculionidae, ship-timber beetles, Lymexylonidae), Hymenoptera (family: horntails or woodwasps, Siricidae), and Lepidoptera (families: leopard moths, Cossidae, and clear-winged moths, Sesiidae). Case stories as to population behavior of stem borers so well known that it needs not to repeat them. It is sufficient to sum up the available data. Self-protection of conductive tissues against stem borers is proceeded with well-expressed CESPPs 2.1.1.1.1.3. "Nonpreference to herbivores, Of unknown nature" and CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent." These CESPPs provide protection of dominants against resident populations of stem borers continually, on condition that operation of them does not fail due to affection by FDESPPs described in the Axiom 8. The operation of these CESPPs forces the resident populations to limit their inhabiting by biomass of the category D - III. Because food resource of D - III is small, an amount of the populations is insufficient to overcome self-protection of D - I and to colonize it. Their density on biomass of the category D-I is Zero. The composition of EEs of stem borers is the same, and they operate in the densityindependent pattern with ctaracteristics Persistent, Potent, and The best. At mass immigration of the bark beetles and long-horned beetles, the level ESPPs 3.1. can decrease to the levels 3.2. or 3.3. All the range of the stem borers concerns the W.C. Cook’s zone (b). 1 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 6.4.2. Life strategy: Openly-feeding defoliators of evergreen coniferous tree species This group of defoliators includes species of the orders Lepidoptera (families: spanworms, Geometridae, lappet moths, Lasiocampidae, tussock moths, Orgyidae, owlet moths, Noctuidae) and Hymenopera (families: conifer sawflies, Diprionidae, leafrolling or webspinning sawflies, Pamphiliidae). Needles of evergreen coniferous species are protected against defoliators by CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent." Needle-eating defoliators present in such ecosystems continually, but if operation of CESPPs is undisturbed, the are forced to limit their feeding by the food stuff having no Antibiosis. The stuff is young needles, in particular on the spruce, where Antibiosis has not developed yet (D-II), staminate “flowers” (D-II), and on weakened needles with decreased self-protection (D – III). Thus, on DI, density of the defoliators is Zero, whereas on D-II and D-III, it is noticeable. Nevertheless, because amount of these categories of biomass is little, in the whole in ecosystems with evergreen coniferous trees as dominants, density of openly feeding defoliators is Insignificant. In the adult stage, the defoliators are able to penetrate in the ecosystems having evergreen coniferous tres species as dominants and lay their eggs. At any numbers of the laid eggs, hatched caterpillars cannot feed by healthy needles, and would die if weakened needles are unavailable. Such suggestion is based on the fail to establish an infestation spot by releasing of larvae of Dendrolimus pini cited in the section 3(1). Further, the absence in some pine ecosystems of perceptible density of Porthetria monacha in the period of an area-wide outbreak of this species suggests that active CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent." is able to suppress an immigration population of the defoliators up to Insignificant level. Again, the immigrants cannot feed everywhere outside of D - III. The above data imply that EE of openly-feeding defoliators of evergreen coniferous tree species on the biomass of the category D - I on the level ESPPs 3.1. "Proper control" provides Zero density with the characteristics Density-independent, Persistent, Potent, The best. It is known the situations, where CESPPs 2.2.1."Natural enemies of invertebrate herbivores" and/or 2.3. "Routine weather suppression" are able to maintain of ESPPs 3.1."Proper control", when an operation of Antibiosis is decreased. Such data concern Dendrolimus sibiricus (an operation of CESPPs 2.3.10.1. and 2.2.1.1.) and D. pini (an operation of CESPPs 2.2.1.). These CESPPs seem to be less reliable for maintenance of ESPPs of the highest level. In so doing, a change of weather situation is able to decrease ESPPs to the level 3.2. "Lag control" or 3.3. "Late control." The composition of EE of openly-feeding defoliators of evergreen coniferous trees is pertinent for the W.C. Cook’s zones (b). In the zones (c) and (d), it takes place modification of the composition caused by the increased role of CESPPs 2.3. “Routine weather suppression.” The zone (a) is unknown for defoliators of the life strategy ’’Openly-feeding defoliators of evergreen coniferous tree species” with the exception of the case 5.4.7. (3.3.), (a) LS(Neodiprion sertifer in young pine plantations) in the Khoper Forest Research and close conditions, where the defoliation exert good effect on vitality overstocked pine monocultures. At mass immigration of the voracious larvae in old instars, the level ESPPs 3.1. can decrease to the levels 3.2. or 3.3. 6.4.3. Life strategy: Bud-mining and web-making defoliators of evergreen coniferous tree species The data concern the single species - the spruce budworm, Choristoneura fumiferana Clem. It needs to consume young foliage of host-trees having decreased Antibiosis. Therefore, in the EE of this species, beside CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent", it operates CESPPs 2.1.2.1.1.3. "Superevasion from herbivores, Exposition to 2.1.1.2.1.2.1." 2 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I Although this defoliator produces vast outbreaks, the level ESPPs 3.1. "Proper control" prevails in ecosystems with dominance of its host-trees. Indeed, the outbreaks are interrupted by decades, when there are no problems with its density. Further, dendrochonologic studies had shown that in past centuries, before structure of forest ecosystems was disturbed by people, outbreaks of the species occurred, but host-trees survived. Therefore, density of the species hardly reached the value High, so that the level of ESPPs decreased to 3.2. “Lag control.” Feeding by staminate "flowers" and young needles, which are free of Antibiosis or have weak Antibiosis allows Choristoneura fumiferana to survive every season. The density of such larvae is Insignificant. Because feeding by staminate “flowers” is common in this species, both these categories of biomass are considered as a whole. Thus, EE of Choristoneura fumiferana on the biomass of the category D - I and staminate “flowers” (D-II) on the level ESPPs 3.1. "Proper control" provides Insignificant density with the characteristics Density-independent, Persistent, Potent, The best. The composition of EE of bud-mining and web-making defoliators of evergreen coniferous trees is pertinent for the W.C. Cook’s zones (b), (c), and (d). In the zone (a), the level ESPPs 3.1. “Proper control” is absent. 6.4.4. Life strategy: Sap-sucking arthropods on buds The buds are protected from sap-sucking arthropods (aphids and mites) by developed CESPPs 2.1.1.2.1.1.1. "Antibiosis to herbivores, Structural, Permanent", and 2.1.1.2.1.2.1. “Antibiosis to herbivores, Physiological (biochemical), Permanent.” In biocenoses, there are no problems with losses of buds due to feeding of sap-sucking arthropods. This differs from the situation in articenoses, where buds of berry crops, in particular the currant, Ribes spp., suffer due to affection by mites. This fact might be explananed by decrease of Antibiosis in buds of crops due to breeding. Thus, in biocenoses, EE of sap-sucking arthropods on buds includes CESPPs 2.1.1.2.1.1.1. and 2.1.1.2.1.2.1., and provides Zero density with the characteristics Density-independent, Persistent, Potent, The best. The W.C. Cook’s zones for sap-sucking arthropods are unknown. 6.4.5. Life strategy: Sap-sucking arthropods on needles, twigs, and shoots Sap-sucking arthropods have traits, which allow them to evade from CESPPs 2.1.1.2.1.2. "Antibiosis to herbivores, Physiological (biochemical)", operating in needles, twigs and shoots. They exert much less negative effect on host-plants than the insects with the chewing mouthparts do. Therefore, it is proposed an operation of CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent", CESPPs 2.1.1.3.1. "Tolerance to herbivores, Deactivation of arthropod digestive juices", and CESPPs 2.2.1. "Natural enemies of invertebrate herbivores." The Antibiosis is weak. It does not kill the arthropods, but decreases its fecundity and the number of generations per season. The density is Insignificant or Low, rarely Intermediate. It operates Directly density-dependent, smooth pattern, Common, Moderate, and The best In the conditions, where activity of natural enemies of invertebrate arthropods is low, in host-plants, it has developed CESPPs 2.1.1.2.1.1.1. "Antibiosis to herbivores, Structural, Permanent", and CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent. The W.C. Cook’s zones for sap-sucking arthropods are unknown. 3 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 6.1.1. BIOCENOSES 6.2.1. The level of ESPPs 3.1. Proper control 6.3.2. The category of biomass D - II An operation of Tolerance to herbivores. Foliage and twigs of deciduous trees and shrubs, needles of the larch, rootlets, reproductive structures (staminate “flowers”, ovaries), seeds in seasons of abundant yielding, regrowth of dominants, tillers and foliage of grassy species, buds as to vertebrate herbivores. The following discource concerns the W.C. Cook’s zone (b). 6.4.7. Defoliators of deciduous tree species of the spring-summer, summer-fall, and fallspring guilds This group includes numerous species of the order Lepidoptera of the spring-summer, summer-fall,and fall-spring guilds (families: lappet moths, Lasiocampidae, and tussock moths, Orgyidae, owlet moths, Noctuidae, tiger moths, Arctiidae, nototontid moths, Notodontidae, closewing moths Pyralidae, and leafrollers, Geometridae, brooshfooted butterflies, Nymphalidae and others). Defoliators of deciduous tree species of the above guilds consume biomass of the category D - II. ESPPs of the level 3.1. "Proper control" as to the above guilds is provided by an operation of CESPPs 2.2.1. "Natural enemies of invertebrate herbivores" and CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues." Being undisturbed, natural enemies keep these defoliators both of resident and immigrated populations on the levels of Insignificant or at least Low densities over unlimited time. As an illustration of the pattern of operation of EEs of these guilds, it might serve the data obtained by the author at studies of Porthetria dispar. The studies were conducted, in oak ecosystems of two categories – in those, where outbreaks of this species were never recorded, and in ecosystems, where the outbreaks had declined not long ago. In the both categories, it was exposed sticky traps with attractive pheromone Porthetria dispar – disparlure. Every such trap caught several males of the species per season year after year. Further, in the similar plots, it was released egg-masses of the species. In some cases, the density of the exposed egg-masses was much more to that capable to excite complete defoliation of the oak. Nevertheless, the number of the caught insects did not increase. Actually all the released insects were killed by their natural enemies. In spite of such a high activity of natural enemies, in the ecosystems, it exists resident populations of the species, although on the Insignificant level of density. EEs of defoliators of deciduous tree species of the spring-summer, summer-fall, and fallspring guilds on the level ESPPs 3.1. "Proper control" provides Insignificant or Low densities. The pattern is Directly density-dependent, smooth, with characterisics Persistent, Potent, The best. 6.4.8. Life strategy: Bud-mining and web-making defoliators of the larch The larch bud moth, Zeiraphera diniana, it seems to be one of a kind in the guild. Its traits are so perfect that the species has been forced out all the competitors in Europe. The information as to situation with this species on the level ESPPs 3.1. "Proper control" is scarce. In the Engadin Valley, within the belt zone, the ecosystem stays as to this species on the level 3.3. "Late control", whereas above and below of the belt zone density of the moth is Insignificant or Low over unlimited period. This fact allows to consider this situation as the level 3.1. "Proper control." EEs of the species in these areas include CESPPs 2.1.2.1.1. "Superevasion from herbivores", 2.3. "Routine weather suppression", and 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant species." 4 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I Thus, the EE of defoliators of the bud-mining and web-making defoliators of the larch on the level ESPPs 3.1. "Proper control" provides Insignificant or Low densities of density-independent pattern with characteristics Common, Potent, The best. 6.4.10. Life strategy: Consumers of seeds having no periodicity of yielding and flowers This group includes numberless species of birds, mammals, insects and mites consuming seeds with a lack of periodicity of yielding. These tree parts belonging to biomass of the category D - II are protected by a cooperation of 2.1.2.3.1. "Supertolerance to herbivores" with CESPPs 2.2."Natural enemies herbivores", 2.2.1. "Natural enemies of invertebrate herbivores" and 2.2.2. "Natural enemies of vertebrate herbivores." In undisturbed ecosystems, there are no problems with regeneration of dominants. The combined effect is sufficient for maintenance changes of generations in dominants. Therefore, the effects are Persistent, Potent, and The best. The available data are insufficient to conclude about patterns of density-dependence in these groups of herbivores. Percentage of seeds of host-trees consumed by herbivores in years of abundant yielding is high. Nevertheless, due to operation of CESPPs 2.1.2.3.1. "Supertolerance to herbivores" and activity of natural enemies in undisturbed ecosystems (CESPPs 2.2.), these losses do not preclude regeneration of dominants. Therefore, the density of the herbivores on the level of ESPPs 3.1. "Proper control" is considered as Intermediate. The density-dependence is unclear. The effect of EE is Common, Moderate, and The best. In the conditions, where consumers of seeds is especially active, it evolved the trait of Antibiosis in seeds. Their density is Low. The pattern is density-independent. The efect is Persistent, Potent, and The best. 6.4.11. Consumers of seeds with periodicity of yielding This group embraces specialized seed-consumers – insects, mammals, for example squirrels, and birds. Again, these plant parts belong to biomass of the category D - II. On them, CESPPs 2.1.2.2.1. "Disappearance from herbivores" operates in a cooperation with 2.1.2.3.1. "Supertolerance to herbivores, and CESPPs 2.2.1. "Natural enemies of invertebrate herbivores", and 2.2.2. “Natural enemies of vertebrate herbivores.” Available data for undisterbed forest ecosystems demonstrate that there are no problems with regeneration of dominants. Therefore, density of these herbivores might be considered as Low. The data concerning the density-dependent pattern operation of this cooperation are insufficient. The effect of EE is Common, Moderate, and The best. In the years of low yielding, the seeds are not protected by any CESPPs , i.e. they belong to biomass of the category D - III. Nevertheless, in this situation, it operates CESPPs 2.5. Therefore, density of the consumers gets decreased that protects seeds on years with abundant yielding. 6.4.12. Life strategy: Vertebrate herbivores of regrowth of dominants in forest ecosystems This group includes the deers, the elks, the roe-bucks, the hares, etc. The regrowth (biomass of the category D - II) is endangered by above animals. It is protected by CESPPs 2.1.2.3.1. "Supertolerance to herbivores." This CESPPs cooperates with CESPPs 2.2.2."Natural enemies of vertebrate predators" and 2.4. "Periodic (bottleneck) suppression." In undisturbed biocenoses, there are problems with regeneration of dominants that gives the grounds to consider density of vertebrate herbivores on the level ESPPs 3.1. "Proper control" as Low. The effect of EE is Persistent, Potent, The best. The pattern of density-dependence is unclear. 6.4.13. Vertebrate herbivores of dominants in grassy ecosystems (Meadow, Steppes, Prairies, Tundra) The experience of savanna reserves in Africa and observations of the pioneers bringing virgin North American prairies under agriculture have shown that natural balance in these ecosystems 5 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I has been kept well. The cooperation of CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues", 2.2.2. "Natural enemies of vertebrate herbivores", and 2.4. "Periodic (bottleneck) suppression" does not allow the herbivores to reach the values of density, which endanger dominants that suggests of presence of continual Low density. This EE operates in the directly density-dependent, smooth pattern and characteristics Persistent, Potent, The best. 6.4.14. Life strategy: Invertebrate chewing herbivores in grassy ecosystems (Meadow, Steppes, Prairies) In self-protection of the dominants prevail CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues." As a cooperator, it serves CESPPs 2.2.1. "Natural enemies of invertebrate herbivores." The advanced Tolerance of host-plants allows the density to reach Low and Intermediate values. The EE operates in directly density-dependent, smooth pattern with characteristics Common, Moderate, The best. Antibiosis in grassy plants against this group of herbivires has developed in specific conditions, where density ot them gets increased. 6.4.6. Life strategy: Sap-sucking arthropods on foliage, tillers and spikes of ecosystems of diverse types Coevolution of sap-sucking arthropods and their host-plants has led to development of CESPPs 2.1.1.3.1.1. "Tolerance to herbivores, Deactivation of arthropod digestive juices", so that the host-plants are able to tolerate the large numbers of these organisms. These traits widen the range of host-plant tissues, which sustain these organisms. That is why they might be considered mainly as consumers of D - II. As cooperators of CESPPs 2.1.1.3.1.1., it serves CESPPs 2.2.1. "Natural enemies of invertebrate herbivores." This composition of EE is common in woody and grassy biocenoses, where numerous species of these organisms present being abundant. Nevertheless, any negative effect on hosttrees is not recorded. Contrary, as it has been shown above, the beneficial effect of them takes place until their density does not reach certain threshold. This EE operates in the directly density-dependent, smooth pattern, characteristics Common, Moderate or Weak, Good. 6.5.3. The W.C. Cook’s zone (c) 6.4.7. Life strategy: Defoliators of deciduous tree species of the spring-summer, summerfall, and fall-spring guilds In the area of the range of a species of defoliators, which defined the W.C. Cook’s zone of possible occurrence (c), a resident populations of herbivore species present continually on the biomass D-II on Insignificant density. At weather situation more favorable for a species, it is known an increase of density of Porthetria distar to the Low level. Due to immigration of individuals from W.C. Cook’s zones more favorable for a species, it is possible appearance of the Low or Intermediate densities over one-two years. CESPPs, which keep resident population on the Insignificant level of density and decrease an immigrating population are 2.3. "Routine weather suppression" and 2.2.1. "Natural enemies of invertebrate herbivores." At an increase of density, it becomes significant the role CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues." The EE operates in the directly density-dependent, smooth pattern with characteristics Common, Potent, The best. 6 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I For the early-spring guild of oak defoliators, the W.C. Cook’s zone (c) is unknown. 6.5.4. The W.C. Cook’s zone (d) 6.4.7. Life strategy: Defoliators of deciduous tree species of the spring-summer, summerfall, and fall-spring guilds In the area of the range of a species of defoliators, which defined as the W.C. Cook’s zone of possible occurrence (d), individuals of this species occurred rarely in as a rule in adult stage. They are immigrants from other W.C. Cook’s zones, which more favorable of existence on them. Here, it operates CESPPs 2.3. "Routine weather suppression." They are so potent that existence of resident populations of defoliators is impossible. Sometimes, these insects are occurred, but they are migrants from outside, which have no chance to reproduce in this area. So that, density of resident population of this species in the zone stays continually on the level Zero. EE is density-independent, Persistent, Potent, The best. Some cases of this category have been cited above at considering of effects of CESPPs 2.3."Routine weather suppression." These are Porthetria dispar in the British Isles, and on the territory situated northwest the Moscow Region (Russia). For the early-spring guild of oak defoliators, the W.C. Cook’s zone (d) is unknown. 6.5. Herbivores in the W.C. Cook’s zones fransformed in the course of coevolution with host- plants Life strategy: the Colorado potato beetle, Leptinotarsa decemlineata and other species The coevolution of the pair "a host-plant – a herbivore" in some circumstances results in extinction of a herbivore within a vast area. The situation of the Colorado potato beetle, Leptinotarsa decemlineata in South America suggests in favor of this view. Why does this species is called the Colorado potato beetle, rather than, for example the Bolivian or Peruvian beetle? The matter of fact, the potato species are of South American origin, and therefore naturally to suggest that the native range of the beetle is situated in this continent. In reality, however, it was unknown in South America, and until new time its range was limited by the Colorado State and the north Mexico. The explanation of this contradiction has been given by P.M. Zhukovsky (1964, pp. 93-94). Considering the cause of absence of Leptinotarsa decemlineata, this scholar supposed that in the ancient time, the ancestry of this species inhabited in South America. However, because climatic conditions of this area were favorable for this species, they resulted in its High density that exerted intensive selective pressure on their host-plants promoting evolving of potato and tomato species with high content of protective substances, in particular demissine and tomatine. Due to insecticide properties of these substances, Leptinotarsa decemlineata has extincted in South America. In fact, all the wild species of the genus Solanum in South America are very rich as to content of the protective substances. The numerous attempts to rear on these species settled larvae of the beetle failed. The quite another situation took place in the range of the beetle in the Colorado State. Severe mountain climate of this area suppressed the beetle to Low density. Therefore, it did not exert significant selective pressure on resident Solanum species. Therefore, they did not evolve the taxa with increased content of protective substances. The situation in South America is a case of conversion of biomass of the category D - II into the category D - I under selective pressure on the part of a defoliator. In a result of this process, at extreme expression of CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent", it arose EE, which provided Zero density of the density-independent pattern, with characteristics Persistent, Potent, The best. From the view of ESPPs, the range of the beetle in South America, where it cannot feed by resident Solanum species, belongs to W.C. Cook’s zone (d), whereas the range in the Colorado 7 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I State belongs to the zones (b) or (c). The areas, where the beetle is a tiresome pest, concerns the zone (a). The extreme expression of the density-independent pattern is a consequence of invading of a predator into an isolated not too large area – an island. A single individual of a predator in such a situation is able to exterminate an endemic species of its prey, which has no protection against this invader. The case of this kind was given by Ch. Elton (1958, Ch. IV). It has been suggested that co-evolution of the pair "a host-plant – a herbivore" in some circumstances leads to extinction of a herbivore in the global scale. Such a suggestion implies that selective pressure on the part herbivores is very potent. This results in developing of their host-plants with high expressed Antibiosis that precludes normal consumption. Just such an explanation has been proposed as a cause of extinction of the dinosaurs at the end of the Chalk geological period (Swain, 1978). According to this hypothesis, intensive consumption of favorable host-plants by the dinosaurs resulted in appearing and spread of plant taxa with high content of alkaloids and tannins. These substances exerted negative effect on reproductive physiology of the animals. The grounds for this suggestion consist in the finding of dinosaur’s eggs with a shell thinner at the end of their era than that in the heyday. The same effect is exerted by DDT of a shell of bird eggs. The extinction of herbivore dinosaurs led to extinction of their predators. That is a destructive effect of very advanced evolution of herbivores in ability to consume of their host-plants. 6.1.1. BIOCENOSES 6.2.1. The level of ESPPs 3.1. Proper control 6.3.3. The category of biomass D - III Absence of operation of CESPPs 2.1. “Plant resistance to PPs”: The annual stem fall and close stuff, foliage of evergreen coniferous species in shadow, and seeds in the years with low yielding. 6.4.1. Life strategy: Stem borers On the level ESPPs 3.1 "Proper control", the ecological niche of stem borers is limited by biomass of the category D - III, including annual stem fall and diverse parts within conductive tissues of healthy trees, with local decreases of Nonpreference and Antibiosis. This limitation is definitely forced, because when the main stock of dominants gets weakened, in any ecosystem, it appeared attacks of stem borers on such trees. This fact implies that inhabiting on the biomass D - III is outside of optimum for stem borers. In it, they suffer due to destructive effects of two kinds. This issue has been discussed in detail in the Section 5(1) as to bark beetles. These speculations might be spread on stem bores as whole. The first is an operation of CESPPs 2.5. "Effects of crowding" characteristic for the springfall period, when the woodpeckers have abundant alternative food, and therefore pursue stem borers slackly. These CESPPs include 2.5.1. "Deterioration and/or shortage of food", 2.5.1.6. "Wounding of insect herbivores at contacts", 2.5.1.6.1. Weakening", 2.5.1.6.1.1. "Mortality due to natural enemies", and 2.5.1.7. "Cannibalism", 2.5.1.7.1. "Direct mortality", and CESPPs 2.5.1.10. Decrease of body weight at shortage of food and/or forced feeding by inadequate food" with the effect 2.5.1.10.1. "Decrease of fecundity and increase of mortality due to weather stress." This EE has the directly density-dependent, smooth pattern. Its characteristics are Persistent, Potent and The best. The latter evaluation of the effect is based on the fact of a decrease of amount of bark beetle populations to the level insufficient for suppression of self-protection of healthy host-trees. In late fall and winter, stem borers on biomass of the category D - III are actively pursued by the woodpeckers. They pertain to CESPPs "Natural enemies of invertebrate herbivores" 2.2.1.2.1. "Vertebrate predators." The pattern of their density-dependence of their EE is rather 8 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I unclear. It might be at shortage of a populations of stem borers, they persue them more forge ahead. This is a sign of the inversely density-dependent patter. It is known cases, when woodpeckers gird heatly trees with the aim to provide themselves with a food – stem borers, which colonize such trees. The potency of their effect is Moderate, because there exist limitations of their impact on stem borers. Some species of stem borers, particularly bark beetles, have traits, which allow them to evade predation by woodpeckers overwintering in the places inaccessible for these birds. In these cases, stem borers are suppressed by exclusively CESPPs 2.5."Effects of crowding." In any case, CESPPs 2.5. "Effects of crowding" operating solely or in a cooperation with 2.2.1.2.1. "Vertebrate predators" keep density of stem borers continually on the values safe for biomass of the category D - I – main stock of dominants. The effect of both of these CESPPs can be characterized as Persistent and The best. 6.4.11. Life strategy: Consumers of seeds with periodicity of yielding (in years with low yielding) In years with low yielding, seeds occur to be of the category biomass D - III. Then, they serve as a trap for seed-consuming insects. The trait of periodicity of seed yielding, i.e. CESPPs 2.1.2.2.1. "Disappearance from herbivore", results in heavy affection of them by invertebrate herbivores when the yield is low. In so doing, seed-consuming insects suffer due to heavy mortality under effect of vertebrate consumers of the seeds particularly the woodpeckers and the squirrels (CESPPs 2.2.1.2.1.). Some species of seed- consuming insects enter in diapause. Thus, it operates CESPPs 2.1.2.2.1. "Disappearance from herbivore", 2.5. "Effects of crowding", in particular 2.5.2. "Attraction of predators and parasites, increase of their searching activity", 2.5.2.1. "Mortality under effect of predators and parasites", and 2.5.1.8. "Diapausation", 2.5.1.8.1. "Mortality under effect of natural enemies." EE is directly density-dependent, smooth, Temporal, Potent, The best. 6.4.2. Life strategy: Openly-feeding defoliators of needles of evergreen coniferous species On the level ESPPs 3.1. "Proper control", resident populations of defoliators of evergreen coniferous species are forced to limit their feeding by weakened needles free from operation of CESPPs 2.1.1.2.1.2.1."Antibiosis to herbivores, Physiological (biochemical), Permanent." This is biomass of the category D - III. Their density stays continually on the Insignificant level. In the case of immigration of the defoliators, their density is suppressed again to the Insignificant level under effect of CESPPs 2.1.1.2.1.2.1. operating in healthy needles. The most probable CESPPs, which suppress density of herbivores on this foodstuff is 2.1.1.2.1.5. “Nutrient inadequacy of host-plant tissues.” The suppressive role of natural enemies (CESPPs 2.2.1.) is probably significant. The EE operates in the is density-independent, Persistent, Potent, The best. 6.1.1. BIOCENOSES 6.2.2. The level of ESPPs 3.2. Lag control 6.3.1. The category of biomass D - I An operation of Antibiosis (and Nonpreference in stem borers) to herbivores. Conductive tissues of stems, branches of trees and shrubs, shoots of coniferous species, roots of the first and the second orders, needles in evergreen coniferous species 6.4.1. Life strategy: Stem borers Let us return to the report by A.A. Berryman and M. Ashraf (1970) about restoring of Antibiosis in Abies grandis trees attacked by Scolytus ventralis at onset of wet weather situation. 9 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I It occurred to be that survivorship of the trees was the less, the more density of the attacked beetles. In this case, the host-trees were weakened (susceptible to the beetles), i.e. ESPPs decreased from the level 3.1. "Proper control." At the colonization by the bark beetles, CESPPs 2.5. "Effects of crowding" did not enter into operation. In fact, it was recorded the resinosis, i.e. CESPPs 2.1.1.2.1.2.2. "Antibiosis to herbivores, Physiological (biochemical), Delayed." Because this CESPPs is of the Extrinsic class, this case indeed is the level ESPPs 3.2 "Lag control." The more density of the attacking beetles, the less their mortality due to resinosis. Therefore, the return on ESPPs of the level 3.1. "Proper control" proceeded at operation of EE in the inversely density-dependent pattern. Because the onset of definite weather situation is a matter of chance, the effect is Spontaneous. Because 5% of the attacked trees die, i.e. the attacking beetles on them survive, this effect is Moderate, and its evaluation is Poor. Depending on value and time of precipitation, the mortality of trees can be less or greater. The latter suggests that the effect would be Bad. 6.4.2. Life strategy: Openly-feeding defoliators of evergreen coniferous species Again, the onset of wet weather situation caused disappearance of young larvae of Dendrolimus pini (V.I. Grimal’s’ky, pers. comm.) and Bupalus piniarius (Klomp, 1966). V.I. Grimal’s’ky showed that the disappearance was a result of increase of oleoresin exudation from needles of the pine. In these cases, the restoration of ESPPs of the level 3.1. "Proper control" took place under CESPPs of the Extrinsic class – CESPPs 2.1.1.2.1.2.2. "Antibiosis to herbivores, Physiological (biochemical), Delayed." The pattern of operation of such EE is seemingly density-independent, Spontaneous and Potent. Because, the young larvae exerted only insignificant damage to their host-trees, the effect should be evaluated as Good. In the case of the greater damage, it can be Poor. 6.1.1. BIOCENOSES 6.2.2. The level of ESPPs 3.2. Lag control 6.3.2. The category of biomass D - II An operation of Tolerance to herbivores. Foliage and twigs of deciduous trees and shrubs, needles of the larch, rootlets, reproductive structures (staminate “flowers”, ovaries), seeds in seasons of abundant yielding, regrowth of dominants, tillers and foliage of grassy species 6.4.3. Life strategy: Bud-mining and web-making defoliators of coniferous evergreen species The outbreak of the spruce budworm, Choristoneura fumiferana Clem. in the Saquenay Region (Quebeck, Canada) has been shown in the Section 5(1) as a special case. 5.4.8 of LS. It demonstrates that ESPPs as to this species can occurs on the level ESPPs 3.2. "Lag control." The EE of this species includes CESPPs 2.1.2.1.1.3. "Superevation from herbivores, Exposition to 2.1.1.2.1.2.1.", CESPPs 2.1.1.2.1.2.1. "Antibiosis to herbivores, Physiological (biochemical), Permanent", and CESPPs 2.1.1.2.1.5.1. "Nutrient inadequacy of host-plant tissues"; on the best part of dominants - 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues." The pattern of operation of the EE is directly density-dependent, delayed. Its effect is Temporal, Potent, and Good. 6.4.7. Defoliators of deciduous tree species of the spring-summer, summer-fall, and fallspring guilds Consider the pattern of density-dependence in the cases of the abrupt cessation of outbreaks of Porthetria dispar at the level of density below than High described above. It took place under the effects of diverse CESPPs, namelly: wandering flocks of insectivorous birds, the increase of activity of parasites or at onset of wet weather situation – cool and prolonged rains. In all these 10 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I cases, it operates CESPPs of the Extrinsic class. Appearing of the birds concerns CESPPs 2.2.1.2.1."Vertebrate predators", the activity of parasites is an operation of CESPPs 2.2.1.1 "Parasites", and onset of the rains is an operation of CESPPs 2.3.3."Cool and prolonged rains in the larval stage of defoliators", 2.3.3.1. "Mortality of larvae at inducing of the acute form of infection." The latter, it operates in a cooperation with CESPPs 2.2.1.3. "Pathogens." 6.4.8. Life strategy: Defoliators of deciduous tree species of the early-spring guild In the defoliators of the early-spring guild, until food resource is not exhausted, density does not effect on mortality. Values of the mortality are determined by weather situation, which in turn effects on time of bud-bursting of host-trees. If the larvae hatch too early, when buds of host-trees still are closed, they die due to starvation. If they hatch too late, when the host-tree leaves have flushed, the larvae are protected badly from natural enemies, and die due to their activity. This is Density-independent pattern, which differs from the above cases by its effect, which is Moderate or Weak. This is so because mortality of the larvae due to a lack of the coincidence of their hatching and the bud-bursting is never complete one. In fact, in East Europe, in the Quercus robur ecosystems, density of the early-spring guild stays never on the Insignificant value. The effect of the EE take place every season. Therefore, it is Persistent. Values of the levels of density are Low, Intermediate or High. Because the density is changed by a chance, this is the level ESPPs 3.2. "Lag control." Unlike to the situation in the level ESPPs 3.3. "Late control", the High density in 3.2."Lag control" stays no more a season. Further, the seasons with High density are not rotated by ones with very decreased density during a number of seasons. Due to advanced capacity in the oak to tolerate losses of foliage at feeding by the early-spring guild, the negative effect of these insects on host-tree vitality is negligible. Therefore, the effect is Good. 6.1.1. BIOCENOSES 6.2.3. The level ESPPs 3.3. Late control 6.3.1. The category of biomass D - I An operation of Antibiosis (and Nonpreference in stem borers) to herbivores. Conductive tissues of stems, branches of trees and shrubs, shoots of coniferous species, roots of the first and the second orders, needles in evergreen coniferous species, sucking arthropods 6.4.1. Life strategy: Stem borers EEs of bark beetles on the level ESPPs 3.3."Late control" have been decribed in detail in the Section 5(1). Here, it is should show differences in EEs of other families of this group. Numerous species of the above familes colonize host-trees jointly. They constitute the same guild. Nevertheless, their habits are different that implies differences in EEs. The longhorned beetles, Cerambycidae are potent competitors in the larval stage. When larvae of bark beetles pupate, larvae of long-horned beetles keep activity, and can consume bark beetle pupae and callow beetles. Full-grown larvae of long-horned beetles pupate in tight "cradles" under bark or in depth of sapwood, where they are inaccessible for subcortial enemies. Contrary, their pupae are favorite food for woodpeckers in fall and winter, when other food of the birds is nearly absent. Having larger body than bark beetles, long-horned beetles spend in preimaginal stages more time. This trait force them to hibernate in sites of development. Here, they are endangered by woodpeckers. In fact, V.I. Gusev (1932) evaluated the mortality of the longhoprned beetle, Monochamus galloprovincialis Oliv. due to predation by woodpeckers as equaled 90%. In adult stage, long-horned beetles are consumed by numerous species of birds. Therefore, CESPPs 2.5.2. "Attraction of preadators and parasites, increase of their searching activity", 2.5.2.1. "Mortality under effect of predators and parasites" is important in EE of longhorned beetles. 11 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I Further, prolonged period of development results in higher demand to quality of food. Probably, this is a serious problem for the long-horned beetles, when resource of weakened hosttrees is close to exhaust. Feeding of them in the adult stage on branches of host trees reflect shortcoming of imaginal feeding. This is operation of CESPPs 2.5.1.10. "Decrease of body weight at shortage of food and/or feeding by inadequate food", 2.5.1.10.1. "Decrease of fecundity and increase of mortality due to weather stress." When adequate food (weakened hosttrees) is exhausts completely, the beetles are forced to oviposite on healthy ones, when the eggs and neonate larvae die in exuding oleoresin or other protective products. This is an operation of CESPPs 2.5.6. "Exhaust of adequate food", 2.5.6.1. "Mortality due to starvation or due to effect of inadequate food in the same generation or next one." The traits of stem borers of the families of Buprestidae, Curculionidae, Siricidae, and the order Lepidoptera are rather similar as to the issue under consideration. Therefore, it is possible to propose EE common for the group of stem borers, except bark beetles. It operates in the Directly density-dependent, spasmodic pattern. Its characteristic is Temporal, Potent, and The bad. This EE is as follows: The level ESPPs 3.3."Late control", a colonized part of biomass D - I, stem borers, except bark beetles, EE - 2.5.2.1.+ 2.5.1.10.1.+ 2.5.6.1., Directly density-dependent, spasmodic, Temporal, Potent, The best. The avaluation of the effect as “The best” is based on the fact that this effect results in decrease of population of the stem borers. Therefore, the population is unable to colonize heathy trees. 6.4.2. Life strategy: Openly-feeding defoliators of evergreen coniferous tree species Outbreaks of defoliators of evergreen coniferous tree species are among most impressive natural hazards. In the species Dendrolimus sibiricus in Siberia, and Porthetria monacha in Europe, the total area of defoliation of dominants and heavy mortality of host-trees during a single outbreak reaches million acres. A decline of these outbreaks is so expressive that is called often a collapse. In EEs of these species on the level ESPPs 3.3. "Late control", it operates numerous subcategories of CESPPs 2.5. "Effects of crowding." The discource in the Section 5 (1) about LSs of defoliators of the group 5.1.1.3.1. "Openly-feeding defoliators of evergreen coniferous tree species" shows that within its range compositions of EEs are inequal, but close ones. Further, all the EEs operate in the Directly density-dependent, spasmodic pattern with characteristics Potent, Temperate, and Bad. 6.4.3. Life strategy: Bud-mining and web-making defoliators of coniferous evergreen species A proposition of EE for this group is based on the single species Choristoneura fumiferana Clem. It includes 2.5.3.1.4. "Mass mortality due to affection by acute form of infection and parasitation", 2.5.1.5.1. "Mortality due to diverse factors", and 2.3.3.1. "Mortality of larvae at inducing of the acute form of infection." The pattern of operation of the EE is Directly densitydependent, spasmodic. Its effects Temporal, Potent, and Bad. 6.1.1. BIOCENOSES 6.2.3. The level of ESPPs 3.3. Late control 6.3.2. The category of biomass D - II An operation of Tolerance to herbivores. Foliage and twigs of deciduous trees, shrubs, needles of the larch, rootlets, reproductive structures (staminate “flowers”, ovaries), seeds in seasons of abundant yielding, regrowth of dominants, tillers and foliage of grassy species 12 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 6.4.9. Life strategy: Bud-mining and web-making defoliators of the larch The EEs of the larch bud moth, Zeiraphera diniana on the level ESPPs 3.3."Late control" in the optimal for this species the belt zone in Engadin Valley was considered in detail in the Section 5(1). To remember, they are the following: EE of Zeiraphera diniana, in the initial outbreak: 2.1.1.3.1.2.+ 2.5.1.1.1.1.+ 2.5.1.2.1.1. +2.5.1.3.1.1.+ +2.5.1.3.2.1.+ 2.5.1.3.2.2.+ +2.5.1.4.1., Directly density-dependent, early spasmodic, Temporal, Potent, Good. EE of Zeiraphera diniana, in the advanced outbreak: 2.1.1.3.1.2.+2.5.3.1.1.+ +2.5.3.1.3.+2.5.3.1.4.+2.5.3.1.5.+2.5.4.1., Directly density-dependent, lately spasmodic, Temporal, Potent, Good, or, if CESPPs 2.1.1.3.1.2. in weakened trees does not operates, - Poor. 6.4.7. Life strategy: Defoliators of deciduous tree species of the spring-summer, summerfall, and fall-spring guilds The EE of the gypsy moth, Porthetria dispar can serve as a model for EEs of the above guilds on the level ESPPs 3.3. "Late control." Our knowledge of population dynamics of this notorious species is lot of much comparing the rest of this group. The EEs of the group include CESPPs 2.5. "Effects of crowding" and CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues." Obviously, parasites and pathogens operate in the Directly density-dependent, spasmodic pattern. Their effect is Temporal and Potent. Due to developed Tolerance of host-trees, significant mortality of them is little of probable. Therefore, the effect should be evaluated as Poor, rather than Bad. The pattern are either early spasmodic or lately spasmodic depending on climate of an area. At the dictincly continental climate, where activity of parasites and pathogens is suppressed, the pattern is lately spasmodic. This is the W.C. Cook’s zone (a). Here, parasites operate weakly as factors of direst mortality and vectors of pathogens. Low humidity of media and high ambient temperatures hinder an operation of the acute form of infection. Affection of a population by pathogens takes place, when this population is overloaded by the slow form of infection. A population stays sevcral years in the conditions of High density until the slow form of infection enters into operation as CESPPs 2.5.3. "Increase of activity of pathogens and parsites in the specific conditions of high host density" with the effect 2.5.3.1.5. "Spontaneous or winter mortality of embryos." In a result, nearly all the embryos die. In the conditions of temperate climate, the term of an arriving of the decline of an outbreak is determined by weather situation. As in the case of Porthetria dispar, drought in the period of the larval stage and severe winter, which suppress entomophagous organisms, which kill insect hosts (preys) and infests them by pathogens. Correspondingly, the decline takes place, when it returns the common weather situation, which favorable for all the natural enemies. The decline is especially dramatic at onset of cool and prolonged rains. Then, nearly all the population dies out over a few days. This is the W.C. Cook’s zone (b). In the areas bordering with the W.C. Cook’s zone (a) and (b), it takes place two stages of the outbreaks – initial and advanced ones. This phenomenon is known for Porthetria dispar in Bashkiria. Due to developed mobility, populations of this species are able to live infestation spots with High density evading from natural enemies. This evasion, however, is not unlimited. Eventually, a population gets overloaded by the slow form of infection, and undergoes a decline. So that, in beginning of an outbreak phase (the initial outbreak) and final of an outbreak phase (the advanced outbreak), it operates diverse EEs. The above conclusions drown for Porthetria dispar might be to some extent be valid for certain other species of the guilds. Thus, the EE of the initial outbreak includes CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues", CESPPs 2.5.2.1. "Mortality under effect of predators and parasites", and CESPPs 2.5.1.4.1. "Mortality due to diverse factors." The pattern 13 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I of this EE is Directly density-dependent, early spasmodic with characteristics Temporal, Potent, and Poor. The EE of the advanced outbreak includes 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues",2.5.3.1.5. "Spontaneous or winter mortality of embryos", 2.3.3.1. "Mortality of larvae at inducing of the acute form of infection", 2.5.3.1.4. "Mass mortality due to affection by acute form of infection and parasitation", and CESPPs 2.5.1.4.1. "Mortality due to diverse factors." The pattern of this EE is Directly densitydependent, lately spasmodic with characteristics Temporal, Potent, and Poor. The predation of insectivorous birds in outbreaks of openly-feeding defoliators is an operation of CESPPs 2.5.2. "Attraction of predators and parasites, increase of their searching activity", 2.5.2.1. "Mortality under effect of predators and parasites." The predation has certain specificity. E.H. Forbush and Ch. H. Fernald (1896, p. 208) expressed it in the following words: "Grimm noticed about Saratov that birds had completely forsaken the places where caterpillars were usually numerous. This is said to have taken place also in the district of Kirsanov in the province of Tambov." Further, these scholars quoted observations by Hermann and Fürst (1893) in Bavaria, which it seemed did not consist with above reports. In fact, in Bavaria, birds aggregated in infestation spot of Porthetria monacha and fed actively by the larvae, pupae and butterflies. These birds were a flock of the starling with about 1000 individuals, the titmice, and the finches. The contradiction might be settles if to take into account that behavior of the birds is determined by different circumstances. At very High density of Porthetria dispar, the nestling suffer probable due to irritation by hairs of older caterpillars of this species. The birds indeed are forced to forsake such forest plots. Here is the report by J.T. Ratzeburg (Tanfil’yev, 1894, p. 137): "Vertebrate animals, especially birds, appear in the beginning of outbreaks of defoliators, when their density is insignificant; but at further reproduction of these insects, when outbreaks spread on vast areas, birds fly out from the areas." The same observations were done by O.A. Grimm (1874, cited in G.I. Tanfil’yev, 1894, p. 137). On the other hand, the cuckoo birds, Cuculus canoris L. can visit affected forest plots for predation on hairy caterpillars. These birds are able to derive intestine of the caterpillars and use it for own consumption. It should recall that the cuckoo birds do not feed own nestlings. Naturally, suppressive effect of the migrating birds is significant in not too large infestation spots, that it was noted else by E.H. Forbush and Ch.H. Fernald (1896, p. 242). Moreover, these scholars cited B. Altum (1880), who reported that a completely defoliated park was used by the titmice, Parus spp. for nesting (Ibid., p. 232). In spring, 20 pairs of the titmice cleared the park from Portheria dispar as whole. The birds fed young caterpillars having no irritating hairs. I.Ya. Shevyriov (cited in G.I. Tanfil’yev, 1894, p. 137) observed intensive feeding of titmice by eggmasses of Porthetria dispar in winter. They, however, were unable to suppress the outbreak. The directly density-dependent, spasmodic pattern of bird’s activity is characteristic for the infestation spots of limited area. There, it is Spontaneous, can be Potent and Good. In the area wide-outbreaks, it takes place the inversely density-dependent pattern of bird’s activity. Its characteristics is Temporal, Weak, and Poor. 6.4.8. Life strategy: Defoliators of deciduous tree species of the early-spring guild In this guild of defoliators, the level ESPPs 3.3. "Late control" takes place in ecosystems with dominance of the English oak, Quercus robur L. at two following situations: a) The undisturbed ecosystems, which most of seasons stay on the level ESPPs 3.2. "Lag control", but in some seasons due to weather fluctuations, density of this guild gets High. They are ecosystems with short term of High density. b) The disturbed ecosystems with long term of High density. These ecosystems have the following characteristics: i) The dominants are composed by diverse phenological forms of the Quercus robur that suppresses operation of CESPPs 2.1.2.1.1. "Superevasion from herbivores." 14 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I ii) The disturbance of ecosystem is so heavy that all the natural enemies of these defoliators are suppressed. iii) The ecosystems are situated in flood plains and are undergo a flooding; this circumstances are favorable for prevalence in the guild Operophthera brumata, because such conditions are perilous for entomophagous organisms of this insects. The EEs of the guld in ecosystems of the categories (a) and (b) are different. The EE of (a) includes CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or . compensation of losses of host-plant tissues" and 2.5.1.5.1 "Mortality due to diverse factors." Its pattern in Directly density-dependent, early spasmodic with characteristics Spontaneous, Potent, and Good. The EE of the point (b) includes CESPPs 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues", 2.5.3.1.5. "Spontaneous or winter mortality of embryos", CESPPs 2.3.1. "Low ambient temperatures at hibernation even they if they are not extraordinary ones" with the effect 2.3.1.1. "Unhatching after hibernation, CESPPs 2.3.6, "Return of frost after thaw" with the effect 2.3.6.1. "Mortality of young larvae in eggs"and CESPPs 2.5.1.5. "Migration on the larval stage with leaving shelters in some species" with the effect 2.5.1.5.1. "Mortality due to diverse factors." Its pattern in Directly density-dependent, lately spasmodic with characteristics Temporal, Potent, and Poor. 6.4.15. Life strategy: Soil-dwelling herbivores in desert and tundra ecosystems The EE of soil-dwelling rodents in thr Desert biome embraces numerous CESPPs: 2.4. ''Periodic (bottle-neck) suppression'' and 2.5. ''Effects of crowding'', when CESPPs 2.4. gets weak. It is probable an operation the following effects of CESPPs 2.5.: 2.5.1. "Deterioration and/or shortage of food", 2.5.1.4. "Emigration in the adult stage in advance of food deterioration and/or exhaust", 2.5.1.4.1. "Mortality due to diverse factors", 2.5.1.10. "Decrease of body weight at shortage of food and/or forced feeding by inafequate food", 2.5.1.10.1. "Decrease of fecundity and increase of mortality due to weather stress", 2.5.2. "Attraction of predators and parasites, increase of their searching activity", 2.5.2.1. "Mortality under effect of predators and parasites", 2.5.3. "Increase of activity of pathogens and parasites in specific conditions of high host density", 2.5.3.1.4 "Mass mortality due to affection by acute form of infection and parasitization", 2.5.4. "Fluctuations of herbivore host resistance to pathogens and parasites as well as virulence of pathogens and aggressiveness of parasites", 2.5.4.1. "Suppression of herbivores over the period, which provides a reprieve for restoring of vitality of dominants", 2.5.5, "Disturbance of media of inhabitation", 2.5.5.1. "Affection by pathogens due to deterioration of sanitary state of the media." This EE operates in Directly density-dependent, spasmodic pattern with chatacteristics Temporal, Potent, and Good. The evaluation "Good" is based of supposition that activity of the rodents is common in desert ecosystems. The EE of soil-dwelling rodents (the lemmings) includes: 2.1.1.3.1.2. "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues", 2.5.1.4. "Emigration in the adult stage in advance of food deterioration and/or exhaust", 2.5.1.4.1. "Mortality due to diverse factors", 2.5.3. "Increase of activity of pathogens and parasites in specific conditions of high host density" 2.5.3.1.4. "Mass mortality due to affection by acute form of infection", 2.5.3.1.6. "Mortality due to inadequate behavior", 2.5.4. "Fluctuations of herbivore host resistance to pathogens and parasites as well as virulence of pathogens and aggressiveness of parasites", 2.5.4.1. "Suppression of herbivores over the period, which provides a reprieve for restoring of vitality of dominants." This EE operates in Directly density-dependent, spasmodic pattern with characteristics Temporal, Potent, and Good. The evaluation "Good" is based of supposition that activity of the rodents is necessary for turnover on substances in tundra ecosystems. 15 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 6.4.5. Life strategy: Sap-sucking arthropods on needles, twigs, and shoots In biocenoses, the level ESPPs 3.3. "Late control" is uncommon for sap-sucking arthropods. The available example of such a situation concerns Adelges (Dreyfusia) picea Ratz., an exotic species, affecting the fir Abies alba Mill. in Europe (Franz, 1956; Karafiat, Franz, 1956; Kloft, 1957). The reports of above scholars have shown that a decline of outbreak of this pest is proceeded by entering into operation of CESPPs 2.5.1. "Deterioration and/or shortage of food", 2.5.1.2. "Raising of secondary structural antibiosis" with the effect 2.5.1.2.1. "Exuding of protective substances on a surface of buds or developing protective barriers in affected tissues" with final effect 2.5.1.2.1.1. "Starvation." This response maintains vitality of affected trees. Therefore, it should be considered as The best. The EE operates in the Directly densitydependent, spasmodic pattern with effects characterized as Persistent, Potent, and The best. 16 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I Table 36. The classification of composition of effective environment (EE) of herbivores of divers groups of life strategies (values of density, patterns of density-dependens, and characteristics of the effects) in biocenoses Levels of ESPPs Category of biomass under protection Density of herbivores in W.C. Cook’s zone 2 3 1 3.1. Proper control 3.1. Proper control 3.1. Proper control Codes of CESPPs operating in EEs of groups of herbivores Patterns of densitydependence 4 5 6 7 8 Density-independent Persistent Potent The best 2.1.1.2.1.2.1., 2.2.1., 2.3. ↔ 6.4.2. Defoliators Density-independent Persistent Potent The best 2.1.1.2.1.2.1., 2.1.2.1.1. ↔ 6.4.3. Defoliators Density-independent Persistent Potent The best Potent The best Zero, (b) 2.1.1.1.1.3., 2.1.1.2.1.2.1. ↔ 6.4.1.Stem borers D - I, needles of evergreen trees Zero, (b), (c), (d) D - I and D – II, needles and staminate “flowers”of evergreen trees Insignificant, (b), (c), and (d) D - I, buds Zero, (b), (c), (d) D - I, needles, twigs, shoots Insignificant, Low, Intermediate, (a), (b) D - II, foliage of deciduous tree species Insignificant or Low , (b); Insignificant (c); Zero (d) D - II, needles of the larch Insignificant or Low D - I, conductive tissues of tree species D - II, seeds having no periodicity of yielding D - II, seeds with periodicity of yielding (in years with abundant yield) Intermediate Low Power of Longevity of Quality of effect of effects of EEs effect of EEs EEs 2.1.1.2.1.1.1., 2.1.1.2.1.2.1. ↔ Density-independent Persistent 6.4.4. Sap-sucking 2.1.1.3.1.1., 2.2.1., ↔ Directly density- dependent, Common 2.1.1.2.1.2.1. 6.4.5.. Sapsmooth sucking 2.2.1., 2.1.1.3.1.2. ↔ 6.4.6. Defoliators Directly density-dependent, Persistent smooth 2.1.2.1.1., 2.2.1.1., 2.3., 2.1.1.3.1.2.. ↔ 6.4.8. Density-independent Defoliators 2.1.2.3.1., 2.2.1., 2.2.2. ↔ Unclear 6.4.10. Consumers of seeds 2.1.2.2.1., 2.1.2.3.1., 2.2.1., 2.2.2. ↔ 6.4.11. Consumers of Unclear seeds Moderate The best Potent The best Common Potent The best Common Moderate The best Common Moderate The best 17 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 2.1.2.3.1., 2.2.2., 2.4. ↔ D - II, regrowth in forest ecosystems Low Persistent Potent The best Persistent 6.4.13. Vertebrate herbivores on smooth dominants in grassy ecosystems 2.1.1.3.1.2., 2.2.1. ↔ 6.4.14. Directly density-dependent, Common Invertebrate herbivores in grassy smooth ecosystems Potent The best ecosystems 2.1.1.3.1.2., 2.2.2., 2.4. ↔ 3.1. Proper control Low D - II in Meadow, Steppe and Prairie ecosystems Low, Intermediate 3.1. Proper control 6.4.12. Vertebrate herbivores on Unclear regrowth of dominants in forest D - II in woody and grassy ecosystems Low, Intermediate Foliage of deciduous tree species, D - II Insignificant, (с) Foliage of deciduous tree Zero, (d) species , D - II D - II, foliage,which obtained The former zone (a) advanced self-protection D - III, conductive tissues tree High, (b) species 3.1. Proper control D - III, seeds with periodicity of yielding (in the years with low yield) D - III, needles of evergreen coniferous species with decreeased antibiosis D – I, conductive tissues with decreased antibiosis 3.2. Lag control D - I, needles of evergreen species with decreased antibiosis High 2.1.1.3.1., 2.2.1., 2.3. ↔ 6.4.6. Sap-sucking Directly density-dependent, Moderate The best Directly-density dependent, Common smooth Moderate Good or Weak 6.4.7. Defoliators Density-independent Common Potent The best 2.3. ↔ 6.4.7. Defoliators Density-independent Persistent Potent The best Density-independent Persistent Potent The best Directly-density dependent, Persistent smooth Potent The best Directly-density dependent, Temporal smooth Potent The best The best 2.3., 2.2.1. ↔ 2.1.1.1.1., 2.1.1.2.1. ↔ 5.3. Defoliators 2.5.1.6.1.1, 2.5.1.7.1.,.2.5.1.10.1., 2.2.1.2.1. ↔ 6.4.1. Stem borers 2.1.2.2.1., 2.5.2.1., 2.5.1.8.1. ↔ 6.4.11. Consumers of seeds Insignificand, (b), (c) 2.1.1.2.1.5., 2.2.1. ↔ 6.4.2. Defoliators Density-independent Persistent Potent Low or Intermediate 2.1.1.2.1.2.2, 2.3. ↔ 6.4.1. Stem borers Inversely densitydependent Spontaneous Moderate Poor or Bad Low, Intermediate 2.1.1.2.1.2.2, 2.3. ↔ 6.4.2. Defoliators Density-independent Spontaneous Potent Good or Poor 18 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I D - II, needles of evergreen species with decreased 3.2. Lag control antibiosis D - II foliage deciduous tree species D - II, foliage of deciduous 3.2. Lag control tree species 3.3. Late control 3.3. Late control Low, Intermediate Low or intermediate Low, Intermediate, sometimes High D - I, conductive tissues of tree species High D - III, conductive tissues of tree species with decreased antibiosis High D - III, needles of evergreen coniferous species with decreased antibiosis D - I and D - II, staminate "flowers" and needles High High 3.3. Late control D - II needles of the larch High 2.1.2.1.1.3., 2.1.1.2.1.2.1, 2.1.1.2.1.5.1, on the best part of dominants - 2.1.1.3.1.2. ↔ 6.4.3. Ch. fumiferana, the special case LS 5.4.8. 2.1.1.3.1.2.,. 2.3., 2.2.1. ↔ 6.4.7. Defoliators 2.1.1.4.1., 2.1.1.3.1.2., 2.2.1. ↔ 6.4.8. Defoliators 2.5.1.6.1.1., 2.5.1.7.1., 2.5.2.1., 2.5.5.2., 2.5.1.10.1. ↔ 6.4.1. Bark beetles on colonized part of D - I 2.5.2.1, 2.5.1.10.1., 2.5.6.1. ↔ 6.4.1. Stem borers, except bark beetles on colonized part of D - I 2.5.2.1., 2.5.3.3., 2.5.3.4., 2.5.1.10.1., 2.1.1.3.1.2.. ↔ 6.4.2. Defoliators 2.5.3.4., 2.3.3.1., 2.5.1.5.1. ↔ 6..4.3 Defoliators 2.1.1.3.1.2., 2.5.1.1.1.1., 2.5.1.2.1.1., 2.5.1.3.1.1., 2.5.1.3.2.1., 2.5.1.3.2.2., 2.5.1.4.1. ↔ 6.4.9. Z. diniana in the initial outbreaks Directly density-dependent, Temporal delayed Potent Directly density-dependent, Spontaneous delayed Moderate Good or Weak Density-independent Temporal Weak Good Directly density-dependent, Temporal spasmodic Potent Bad Directly density-dependent, Temporal spasmodic Potent The best Directly density-dependent, Temporal spasmodic Potent Bad Directly density-dependent, Temporal spasmodic Potent Bad Directly density-dependent, Temporal early spasmodic Potent Good Good D - II needles of the larch High 2.1.1.3.1.2., 2.5.3.1., 2.5.3.3., 2.5.3.4., 2.5.3.5., 2.5.4.1. ↔ 6.4.9. Z. diniana in the advanced outbreaks Temporal Potent Good, or if 2.1.1.3.1.2. in weakened trees does not operate, - Poor D - II, foliage of deciduous tree species High 2.1.1.3.1.2., 2.5.2.1., Directly density-dependent 2.5.1.4.1. ↔ 6.4.7. Temporal early spasmodic Defoliators in initial outbreak Potent Poor 3.3. Late control Directly density-dependen, lately spasmodic 19 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I High D - II, foliage of deciduous 3.3. Late control tree species High High 3.3. Late control 3.3. Late control D - II, foliage of deciduous tree species High D – II, grassy vegetation in Desert ecosystems High D – II, grassy vegetation in Tundra ecosystems High D - I and D - II High 2.1.1.3.1.2., 2.5.3.5., 2.3.3.1., 2.5.3.4., 2.5.1.4.1. ↔ 6.4.7. Defoliators in advanced outbreak 2.5.2.1. Avian predators ↔ 6.4.7. Defoliators in small infestation spots 2.5. 2.1. Avian predators ↔ 6.4.7. Defoliators in area-wide outbreaks 2.1.1.3.1.2., 2.5.1.5.1. ↔ 6.4.8. Defoliators at short term of High density 2.1.1.3.1.2., 2.5.3.5., 2.3.1.1., 2.3.6.1., 2.5.1.5.1. ↔ 6.4.8. Defoliators at long term of High density 2.5.1.4.1., 2.5.1.10.1., 2.5.2.1., 2.5.3.4., 2.5.4.1., 2.5.5.1. ↔ 6.4.15. Soil-dwelling herbivores in desert 2.1.1.3.1.2., 2.5.1.4.1., 2.5.3.4., 2.5.3.6., and 2.5.4.1. ↔ 6.4.15. Soil- dwelling herbivores in tundra 2.5.1.2.1.1. ↔ 6.4.5. Sapsucking arthropods Directly density-dependent, Temporal lately spasmodic Potent Poor Directly density-dependent, Spontaneous spasmodic Potent Good Inversely densitydependent, in area-wide outbreaks Weak Poor Directly density-dependent, Spontaneous early spasmodic Potent Good Directly density-dependent, Temporal lately spasmodic Potent Poor Directly density-dependent, Temporal spasmodic Potent Good Directly density-dependent, Temporal spasmodic Potent Good Directly density-dependent, Temporal spasmodic Potent The best Temporal 20 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 6.1.2. ARTICENOSES Below, it will be considered EEs of herbivores at operation of CESPPs 2.6. "Human control measures", which are known to be effective for maintenance of the level of ESPPs A.3.1. “Proper control” for herbivores and phytopathogens. The data are presented in the Table 37. Table 37. The classification of composition of effective environment (EE) of herbivores of diverse groups of life strategies (values of density, patterns of density-dependens, and characteristics of the effects) at operation of CESPPs 2.6. "Human control measures" in articenoses based on the cases of successful usage of the measures on the level ESPPs A.3.1.”Proper control.” Codes of CESPPs 2.6. (Antropic class) 1 2.6.1.1.1. 2.6.1.1.2. 2.6.1.2. 2.6.1.2. . 2.6.1.3. Category of biomass under protection 2 D - I, buds and shoots of pines D - II, Stems of the wheat and foliage and stems of potato D - II, foliage and stems of the winter wheat and the potato D - I, all the parts of pines D - II, stems and foliage of the winter wheat and the potato D - I and D - Il, conductive and assimilative tissues Codes of CESPPs 2.1., 2.2., 2,3., Density of and 2.4. (Extrinsic class) using by herbivores and people for suppression of PPs and affection by measures invented by people phytopathogens (species and groups of PPs are noted) 3 4 Zero Zero Zero Insignificant or Low Zero Zero 2.1.1.2.1. Evetria (Petrova) spp. Patterns of densitydependence 5 Density-independent 2.1.1.2.1. Mayetiola destructor on winter wheat; Leptinotarsa Density-independent decemlineata on the potato A.2.1.2.1.1, A.2.1.1.3.1.2, A.2.1.1.2.1.1. herbivores on the winter wheat Inversely density-dependent A.2.1.2.1., A.2.1.1.3.1.2. on the potato 2.1.1.2.1., 2.1.1.2.2. Stem borers, Density-independent defoliators and phytopathogens Longevity Power Quality of effects of effect of effect of EEs of EEs of EEs 6 7 8 Persistent Potent The best Persistent Potent The best Persistent Moderate The best Persistent Potent The best Low or Insignificant A.2.1.1.3.1., A.2.1.1.4.1., A.2.1.1.2.1. Invertebrate herbivores Inversely density -dependent Persistent Moderate Good Insignificant 2.2. Defoliators and sap-sucking arthropods Directly density- dependent, smooth Persistent Potent The best 21 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 2.6.1.4. 2.6.1.5. 2.6.1.6. 2.6.2.1. 2.6.2.2. 2.6.2.3. 2.6.3.1. 2.6.3.2. 2.6.3.3. D - II, Foliage or conductive tissues of deciduous tree species, seeds D - II foliage of the potato D - I and D - II, all the plant parts in glasshouse culture D - I and D - II D - I and D - II D - I and D - II, all the plant parts D - I and D - II, all the plant parts D - I and D - II, all the plant parts D - I and D - II, all the plant parts Low or Intermediate 2.1.1.3.1., Defoliators and sap-sucking arthropods Directly density-dependent, smooth or delayed Persistent Potent of Good Moderate Insignificant or Zero 2.3. Aphids – vectors of viruses Density-independent Persistent Potent The best Insignificant or Zero 2.4. All the PPs of a given crop Density-independent Temporal Potent The best Density-independent Persistent Potent The best Inversely density-dependent Persistent Density-independent Persistent Density-independent or inversely density-dependent Temporal Zero Insignificant or Zero Insignificant or Zero Low or Intermediate Eradication of burdensome invaded herbivores Eradication of burdensome resident herbivores Eradication of burdensome phytopathogens or herbivores A.2.1.1.4. and control all the complex of herbivores Insignificant or Zero Control of a purpose herbivore species Density-independent Temporal Insignificant Control of a purpose herbivore species Directly density-dependent, smooth Temporal Potent or Moderate Potent or Moderate Potent of Moderate Potent or Moderate Potent The best The best Good Good The best 22 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I CESPPs 2.6.1.1. Growing of cultivars resistant to plant pests 2.6.1.1.1. Growing of plant taxa with natural resistance of CESPPs 2.1. In pine plantations established in south Ukraine, it is almost impossible to find the pine shoot moths, Evetria (Rhyacionia, Petrova) spp. on the Crimean pine, Pinus pallasiana, although the moths affect heavily the Scots pine, Pinus sylvestris even if the former grows as an admixture in the stands with the latter. Hence, P. pallasiana keeps resistance in spite of High density of the pest insects. Moreover, the insects lay their eggs on the buds of P. pallasiana, but all the larvae die due to 2.1.1.2.1.2. "Antibiosis to herbivores, Physiological (biochemical)." This is the case of operation CESPPs 2.6.1.1.1. "Growing of cultivars resistant to plant pests" providing Zero density with the density-independent pattern, Persistent, Potent, and The best. 2.6.1.1.2. Growing of cultivars bred with CESPPs A.2.1. In Kanzas, at growing of varieties of the wheat resistant to Mayetiola destructor, it was almost impossible to find any evidence of this insect at weather situation unfavorable for it. Again, varieties of the potato with developed Antibiosis actually free from damage due to the Colorado potato beetle, Leptinotarsa decemlineata in spite of amount of populations of this species is continually great. In both above cases, the method 2.6.1.1.1. "Usage of CESPPs of the category 2.1." exploits CESPPs A.2.1.1.2.1. "Antibiosis to herbivores" The method operates in the density-independent pattern. Its effects are Persistent and Potent. From the view of suppression of the beetle, this effect should be evaluated as The best. However, from other views the evaluation may be negative. In particular, the potato varieties with developed Antibiosis against Leptinotarsa decemlineata bred in USSR cannot be used for the food purpose due to inadmissibly high content of toxic substances in tubers. In Ukraine, in the Forest-Steppe biome, it has been bred and used in agricultural practice great number of winter wheat varieties with well-developed traits serving as CESPPs A.2.1.1.3.1.2 "Tolerance to herbivores, Repair or compensation of losses of host-plant tissues" and A.2.1.2.1.1. "Superevasion from herbivores" against a number of species of stem flies, sapsucking arthropods, defoliators, and root insect pests. Against the stem sawfly, Cephus pygmaeus, it operates A.2.1.1.2.1.1.1. "Antibiosis to herbivores, Structural, Permanent." In the conditions, where the author’s studies were conducted, these CESPPs occurred to sufficient for reliable protection against all the complex arthropod pests over the period of studies – 15 years. The density-dependence in this case is the inversely density-dependent one. This is so because the method would be effective on condition that amount of pest populations is not too much. The keeping of pest populations on the safe for the winter wheat level is exerted by climate of the area, i.e. CESPPs 2.3. "Routine weather suppression", 2.4. "Periodic (bottle-neck) suppression", and 2.2.1. "Natural enemies of invertebrate herbivores." If these CESPPs would be relaxed, an amount of herbivore populations would be great. In such situation, CESPPs concerned plant resistance to herbivores are invalid. It was recorded small infestation spots of some insect pests, but they quickly declined obviously due to operation of the above CESPPs. This is so because activity of the cooperators of as a whole is very significant. The effect of the pattern is Persistent. It is Moderate, because the self-protection of the host-plant is only one of many CESPPs, which determine the lack of damage on the part of the herbivores. In given conditions, the effect is The best. 2.6.1.2. The usage of cultural practices, which maintain ESPPs by keeping healthy state of plants In plantations of evergreen coniferous tree species, it is used wild species, which possess well-developed CESPPs 2.1.1.1.1. "Nonpreference to herbivores" and 2.1.1.2.1. "Antibiosis to herbivores." Therefore, for maintenance of ESPPs of the level A. 3.1. "Proper control" in these articenoses, it is sufficient to choose tree species adapted to local conditions (CESPPs 2.6.1.1.1.) 23 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I and to grow them at silvicultural practices, which provide an operation of the prerequisite of CESPPs 2.1.1.1.1. "Nonpreference to herbivores" and 2.1.1.2.1. "Antibiosis to herbivores." This is a proper physiological state of the trees. Just this is provided as to all the PPs by CESPPs 2.6.1.2. The cases of the level of ESPPs 3.1. "Proper control" in plantations of Pinus pallasiana and P. sylvestris show that this aim is achieved at the sound management. Because this CESPPs provides the complete lack of the damage by PPs, it operates in the density-independent pattern. Its effects are Persistent, Potent, and The best. Complex of cultural practices at growing of the winter wheat maintains the level of ESPPs A.3.1. "Proper control" providing host-plants by prerequisites of CESPPs. Because this method is used as to the category of biomass D - II, its efficacy is dependent on activity of cooperators from the part of diverse CESPPs. When the cooperators weaken in their activity, the amount of herbivore populations increases, and CESPPs 2.6.1.2. fails. Hence, this CESPPs operates in the inversely density-dependent pattern. In the conditions of the region, where the author’s studied were conducted (the Forest-Steppe biome of Ukraine), its effects are Persistent, Moderate, and The best. 2.6.1.3. Introducing of natural enemies of herbivores At introducing of natural enemies of herbivores, it should reach two aims: they need to suppress their preys to the level below threshold of damage of plants, but do not kill the preys completely. In the opposite case, the introduced natural enemies would extinct if they are monophagous. Hence, it should takes place directly density-dependent, smooth pattern. It is known the case, when invaded by chance a parasite wiped out its host in some ecosystems and after that disappeared. The host, however, survived in other places, penetrated into the ecosystems, where the parasite was absent, and reached High density. Such case was reported by T.H.C. Taylor (1937) for the beetle, Promecotheca reichei and the parasitic mite, Pediculoides ventricosus in Fiji. Thus, to be successful, introducing natural enemies should operate in the directly densitydependent, smooth pattern. Its effect must be Persistent, Potent. Then, it is The best. In articenoses, where activity of predators is low, sap-sucking arthropods often get serious pests. In this context, consider the case of the successful suppression of the exotic coccid, Icerya purchasi Mask. by the introducing of the lady beetle, Rodolia cardinalis Muls. This case concerns the present discourse, because it simulates the situation in biocenoses. H.L. Sweetman (1958, Ch. IX) in his review reported that in a result of the introduction, the coccid is usually kept on Insinificant density, i.e. below threshold of damage of host-trees. This is the directly density-dependent, smooth pattern with the effect Persistent, Potent, and The best. In some orchards, the prey gets disappears completely (Ibid.). This is density-independent pattern, which is of a doubtful value, because it would results in an extinction of the monophagous predator, and fluctuation of pest density. 2.6.1.4. Promotion to native natural enemies of herbivores Resident natural enemies of herbivores, when conditions of their existence are favorable, suppress their hosts of prey in the directly density-dependent, smooth pattern. Its effects are Persistent, Potent, and The best. At cultural practices for promotion to them, it should aim just the same situation. 2.6.1.5. Cultivation of plants in climatic conditions that suppress plant pests This CESPPs effective as to maintain of ESPPs on the level A.3.1. "Proper control" if it operates in the density-independent pattern, when subject PPs either are kept continually on Zero density or Insignificant one. Thus, CESPPs 2.6.1.5. operate in the density-independent pattern. Its effects should be Persistent, Potent, and The best. 24 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 2.6.1.6. Simulation of the conditions analogous an operation of CESPPs 2.4. Periodic (bottle-neck) suppression This CESPPs again is directed on complete destruction of PPs. Hence, the pattern of its operation is density-independent with the effects Persistent, Potent, and The best. 2.6.2. Artificial measures of long-lasting character 2.6.2.1. Eradication of plant pests by mechanical means This CESPPs is very expensive, and it justifies expenses if an invaded PPs is destroyed up to last individual. As an evidence of a need so radical destroying, it serves the case of invasion on of the western rootworm, Diabrotica virgifera virgifera LeConte in Europe, which bears threaten growing of the corn over the continent, has begun from a single gravid female transported in a plane from Chicago to Belgrad in 1980-ies (R. Edwards, pers. comm.). It has been known the cases of success and failures in these enterprises. It concerns Leptinotarsa decemlineata in Europe. This pest penetrated from America to Europe several times. S.P. Ivanov et al. (1938, p. 22) reported that it was found out in Britain and Germany in 1876, 1881, 1887 and 1914. In all the cases, the invaders were destroyed completely. But in 1920, it was penetrated into France. On this occasion, it was not wiped out, and eventually firmly established in Eurasia. The above cases implies, that the CESPPs 2.6.3.1.1. is an extreme expression of the densityindependent pattern directed on complete eradication of PPs. If it takes place, its effects are Persistent, Potent, and The best. 2.6.2.2. Eradication of plant pests by release of sterilized individuals or breakage of pheromonal communication The eradication of a herbivore species by releasing of sterilized individuals is reliable on condition that a resident population of the same species has been decreased to very Low density by temporary suppressive measures. Then, if the amount of sterilized individuals exceeds greatly the amount of a resident population, the suppression over an unlimited period gets probable. Of course, this is achieved on limited areas, especially on islands. The breakage of pheromonal communication is based on the similar principle. For example, this is the planning of suppression of Porthetria dispar by M. Berosa et al. (1974, cited in H.C. Coppel and J.W. Mertins, 1977). An island population of this species should be decreased by application with insecticide, after that is used exposition of traps with disparlure, and/or application of it with microcapsulated disparlure. Obviously, this method has inversely density-dependent pattern. In the case of the success, its effects are Persistent, Potent, and The best. 2.6.2.3. Change of cultivated crops In all the case, the pattern of this CESPPs is density-independent, but potency of it can be varied. When the affected crop is deleted completely on a given area, the effects are Persistent, Potent, and The best. If the activity is limited by diversification of growing crops an affected one is used in shift of crops, the a risk of the damage to some extent can be continued. In this case, the level of potency of the effect is Moderate. 25 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I 6.1.2. ARTICENOSES 6.1.2.1. The level ESPPs 6.1.2.1.2. The level of ESPPs A.3.2. Lag control. 2.6.3. Temporary suppressive measures 2.6.3.1. Cultural practices with the aim to control plant pests They operate in the density-independent pattern. Their effects are Temporal, Moderate or Potent, and The best. 2.6.3.2. Sanitary measures They operate again in the density-independent pattern. Their effects are Temporal, Moderate or Potent, and The best. 2.6.3.3. Temporary suppressive measures sensu stricta. A. McFadyen (1963, Ch. XVIII) supposes that effect of application of insecticides does not depend on density of a herbivore population, i.e. it does not excite greater mortality in populations with High density. This is not true. Toxic effect of insecticides depends of physiological state of a herbivore population. The state is worst in overcrowded populations. Knowledge of this fact allows usage less rates of insecticides combining them with microbiological preparations. The latter decreases physiological state of a herbivore population, so that an insecticide with a lower rate of application exerts the same mortality as that at a full rate. CESPPs 2.6.2.3. operates in the directly density-dependent, smooth pattern. Its effects are Potent, because density decreases to Insignificant level, Temporal because the decrease continues only a single generation of a pest insect. In articenoses, particularly in agriculture, in the most cases, CESPPs 2.6.3.3. is the only means, which is able to save an amount of yield and/or its quality with high justification of expenses. In biocenoses, application of often is necessary for saving of dominants or for maintenance of their concern. At the sound application, of insecticides bear a minimal danger for people and the environment. Therefore, this CESPPs might be evaluated as The best. 6.1.1. BIOCENOSES and 6.1.2. ARTICENOSES The factors providing the conservation of herbivore populations on the safe for dominants density in biocenoses and articenoses To exist, ecosystems must maintain an operation of two processes of the opposite character. The first is keeping of PPs on the level below threshold of damage for dominants. The second process is a conservation of them on safe level of density that is necessary for turnover of substances in ecosystems. Consider the ways of operation of the second process in diverse groups of herbivores. Stem borers The continual presence of stem borers in forest ecosystems might be explained easily. They thrive on the level ESPPs 3.1. "Proper control", because every year some part of the biomass of the category D - I transforms into D - III. In the given case, the part is presented by conductive tissues of trees. This is a waste part of dominants, which is not protected any CESPPs of the categories 2.1., 2.3., and 2.4. Here, it operates CESPPs 2.2.1. "Natural enemies of invertebrate herbivores, 2.2.1.2. "Vertebrate predators" and the Intrinsic class – 2.5. "Effects of crowding." Among natural enemies of stem borers, most important are the woodpeckers (CESPPs of the category 2.2.1.2.1.). Nevertheless, there exist a number of limitations, which allow stem borers 26 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I to survive in spite of predation by the woodpeckers. This issue has been considered in the Section 5(1). Bud-mining and web-making defoliators On the levels Insignificant, Low and Intermediate densities, these insects are relatively wellprotected by web cover against all the natural enemies. In the imaginal stage, the are protected by mimicry, as Tortrix viridana, or by fly in the period of low activity of avian predators, as Operophthera brumata. Their eggs are laid so perfectly that they nearly inaccessible for parasites and predators. Therefore, the species consuming foliage of deciduous trees are numerous every year except the innocuous phase on the level ESPPs 3.3. "Late control." Here, at a decline of outbreaks, a population undergoes deep depression over a number of years. Openly-feeding defoliators At Insignificant level of density, defoliators become slight for parasites and predators. The effect of density on level of parasitization was proved by a number of experiments (the review has been offered by G. Morrison and D.R. Strong, 1980). In particular, in studies by T. Burnett (1958), the parasite wasp, Encarsia formosa Gahan in glasshouses parasititized 40% of its host – the whitefly, Aleyrodes vaporariorum Westw. at the lowest density and 80% or more at high values of the density. Staying in shelters, defoliators suffer due to parasites also depending on density. For example, the parasite Exenterus abruptorius affects the European pine sawfly, Neodiprion sertifer in forest litter beginning with its density two-three cocoons per square meter of the soil surface (Gur’yanova, 1984a). As to mammal ground predators, the similar dependence was shown again for Neodiprion sertifer (Holling, 1963). The same is known for avian predators, which concentrate their attention on the most numerous species of preys, whereas widely sparse insects have a good chance to be unnoticed (Tinbergen, 1960; Korol’kova, 1963, p. 63). Some species of defoliators are protected from birds by mimicry, other have poisonous hairy cover, and such as Porthetria dispar spend day-hours, when the bird are active, in shelters and pupate in these shelters. Thus, the phenomenon "specific searching image" is characteristic for diverse interrelations of herbivores with entomophagous organisms. The less attacks on the part of stinging parasites – vectors of infection, the greater health of a population, because individuals with high-virulent strains of pathogens died out with their hosts being impossible to infest the rest of a population. Healthy caterpillars are able to repulse attacking parasites throw off them by strokes of their body. Some part of individuals within a common insect host rejected by parasites at oviposition. At last, the oviposition proceeds successfully, but parasite’s eggs in body of healthy hosts meet with protective response of the immune system. The eggs get safe for the hosts. The phenomenon of insect host’s resistance to parasites deserves considering of the case stories. N. F. Meier (1921, 1926, 1929, cited in S.P. Ivanov et al., 1938, p. 45-46) conducted the studies on exposition of the cabbage moth, Plutella maculipennis Curt. by its parasite wasps Angitia fenestris Holmgr., Exetastes cinctipes Ratz., Anilasta ebenina Crav., Apanteles glomeratus L., and others. It was found out that 15% of larvae of this moth were rejected by the parasites, although some other larvae of this species were parasitized repeatedly. In similar tests of the same scholar with the moth, Pieris brassicae L., up to 40% of the parasitized larvae killed laid eggs by means the response of phagocytosis and survived to adults. This was shown for the parasites Anilasta ebenina and Apanteles glomeratus. As to protection of insect hosts from parasites by means of phagosytosis, S.P. Ivanov et al., 1938, p. 45) quoted also F. Tolg (1910), Smith van Burst (1919), and Fr. Eckstein (1931). Another means of self-protection of insect host is encapsulation of parasite’s eggs (Van den Bosch, 1964). 27 G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH. PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I Against tachinid parasites laying eggs on a surface of larvae, it serves a throwing off the eggs with the larval skin at molting (Ivanov et al., 1938, p. 46). When laying their eggs, females of defoliators use some dodges to protect them against parasites. For example, the apple tent moth, Hyponomeuta malinelus produces egg-masses in several layers. The eggs in lower layers are inaccessible for the highly specific parasite Ageniaspis fuscicollis Dalm. In a result, up to 30% of the eggs survive (M.M. Padiy, pers. comm.). Sap-sucking arthropods The sap-sucking arthropods inform us about one more means how herbivores survive in spite of high efficiency of a predator. The case is the interrelations between the exotic coccid, Icerya purchasi and its introduced predator - the lady beetle, Rodolia cardinalis in the conditions of the Caucasian Black Sea coast. Here, the lady beetle occurred to be very effective, so that damage on the part of the coccid was stopped. Nevertheless, the coccid exists at the noticeable density providing its predator by food. B.I. Rukavishnikov (1964, p. 289) in comments to the Russian translation of the book by H.L. Sweetman offered data of E.M. Stepanov (1935), who showed some circumstances that allowed the coccid to survive on Low density. Little infestations of the coccid were located on wild leguminous species with high content of alkaloids, for example on the genera Spartium and Genista. The lady beetle denied feeding by this prey due to presence alkaloids in its body. These plants had no economic value, so that the presence of the coccid was safe. From the infestations, the coccid spread into orchards. Further, the coccids entered into symbiosis with resident species of ants, which served as their bodyguards. The ants were attracted by sweet exudates of the coccids, and built cameras from clay for their protection. If the lady beetles aimed to penetrate into the cameras, the ants instantly expelled them. In this story, it is notably that capacity to consume the plants with developed Antibiosis gives a herbivore resistance to a predator. This is a situation concerned the triotrophic interactions – a concept established by K.J. Stark et al. (1972). According to the concept, a state of host-plants effects on traits of their consumers, and the latter determines affection of the consumers by natural enemies. Again, such relations were traced in the plant-aphid-lady beetle system (Giles et al., 2002). Carpophagous herbivores The triotrophic interactions might be found out in this group of consumers on the base of studies by G.A. Viktorov (1967, p. 29). He noted that seeds of the wild grassy leguminous species were affected very little by the moth, Etiella zinckenella Tr., and explained this fact by high activity of its parasites. Contrary, seeds of wild woody species Robinia pseudoacacia and Caragana arborescence were affected heavily. The explanation of this difference consists in the fact that the parasites Bracon piger and Epiurus ventricosa cannot pierce solid walls of pods of the woody species by their ovipositors. As to the parasite Limneria fuscicarpus Thomp., the larvae fed on the woody species occurred to be immune to this parasite. In this case protective substances of host-plants served for protection of their consumers from parasites. Notably that in the grassy leguminous species, it operated CESPPs 2.1.2.3.1. "Superolerance to herbivores" in a cooperation with CESPPs 2.2.1. "Natural enemies of invertebrate herbivores, 2.2.1.1. "Parasites", which offered more reliable protection of seeds than CESPPs 2.1.1.2.1. "Antibiosis to herbivores" (in the woody plants). 28