Ebook "STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH"
Gleb I.Vasechko,
Cand.Sc.(Biology)
© Gleb Vasechko, 2005 All rights reserved.
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Created 20.03.2005
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Gleb Vasechko.
PART I. A LIST OF THE AXIOMS. Continuation.
AXIOM 8. FACTORS of DISTURBANCE of ECOSYSTEM STABILITY TO PLANT PESTS and THEIR EFFECTS
ESPPs to the wide range of PPs in biocenoses is known to stay on the level 3.1. "Proper control " over unlimited time. In articenoses, the level ESPPs A.3.1. "Proper control " is also possible. The opposite situations when ESPPs decreases to the levels 3.2. "Lag control " or 3.3. "Late control " are known even more well. The lowering of ESPPs from the level 3.1. to the levels 3.2. and 3.3. means an affection of ecosystems by PPs.
The capacity to stay on the level 3.1. "Proper control " is immanent for biocenoses. This is a result of the Darwinian natural selection. On the other hand, the lowering from this level is a result of stresses, which in the given text will be considered as factors of disturbance of ESPPs (FDESPPs). As to articenoses, the level ESPPs A.3.1. "Proper control " is maintained by artificial selection and cultural practices, whereas the lowering from the level ESPPs A.3.1. "Proper control " is again a result of operation of FDESPPs.
The Axiom 8 declares known FDESPPs, classifies them anddemonstrates their effects - the character of affection of dominants by PPs (Table 8).
The first step of the classification (8.1. - 8.6.) is subdivision of the wide range of FDESPPs on the categories according to their capacity to break of prerequisites of CESPPs. Such categories are the following:
8.1. "Anthropic-1. They break prerequisites A.2.1.1.P.1., A.2.1.2.P.2.., 2.2.1.P.2., 2.2.1.P.9., 2.3.P.2. and 2.4.P.1. due to human activity.
8.2. "Anthropic-2. They break prerequisites 2.1.1.P.2., A.2.1.1.P.2., 2.1.2.P.1., A.2.1.2.P.2, A.2.1.2.4.1., 2.2.1.P.3. -.2.2.1.P.9., and 2.2.2.P.1. -2.2.2.P.3. due to human activity.
8.3. "Weather disturbance. " They break prerequisites 2.3.P.1., 2.3.P.2., 2.3.P.3., and 2.3.P.4.
8.4. "Endogenic disturbance. " They break prerequisites 2.1.1.P.2., 2.1.1.P.1., A..2.1.1.P.1., and 2.5.P.2.
8.5. "Force-majeure anthropic impacts. "
8.6. "Force-majeure natural impacts. "
The second step of the classification (8.1.1. - 8.6.5.) clarifies the substance of the FDESPPs, which exert the failure of the prerequisites of CESPPs.
The third step of classification (8.1.1.1. - 8.6.5.1.) shows the objects, on which the above FDESPPs are directed. They are CESPPs, which failed to operate due to affect of a given category of FDESPPs.
The forth step of the classification (8.1.1.1.1. - 8.6.5.1.1.) shows the terminal effect of activity of FDESPPs -a character of affection of dominants by PPs.
At last, the fifth step of the classification shows FDESPPs cooperating usually with a given FDESPPs.
The short characteristic of FDESPPs and their objects are the following.
The category 8.1. "Anthropic-1 (breakage of prerequisites A.2.1.1.P.1., A.2.1.2.4.1., 2.2.1.P.1., and 2.2.1.P.2. due to human activity). " This means that in result of breeding, dominants unable to express self-protection against PPs in the certain conditions where they would be grown. The capacity of self-protection absents in its genotype being lost in a process of breeding.
The category 8.2. "Anthropic-2 (breakage of prerequisites 2.1.1.P.2., A.2.1.1.P.2., 2.1.2.P.1., A.2.1.2.P.2, 2.2.1.P.3. -.2.2.1.P.9., and 2.2.2.P.1. -2.2.2.P.3. due to human activity) " concerns the cultural practices in ecosystems (biocenoses and articenoses), and other human activities, which threaten ecosystems.
The category 8.3. "Weather disturbances. " (breakage of prerequisites 2.3.P.1., 2.3.P.2., 2.3.P.3., and 2.3.P.4. ) embraces the firstly, effects of weather situations with significant deviations from yearly average indices for a given territory, including heavy droughts, severe winters (air temperatures close to minimal ones at weak snow cover, sharp decreases of air temperatures in a season). Such stressors have a destructive effect both on physiological state of plants, and activity of natural enemies of PPs, i.e. FDESPPs 8.3 cause failure of operation of CESPPs 2.1. "Plant resistance to PPs " an 2.2.1. "Natural enemies of invertebrate herbivores. "
Also, 8.3. "Weather disturbances, embraces also the effects of specific weather situations, which promote diverse phenomena of vitality of PPs. This is a failure of operation of CESPPs 2.3. "Routine weather suppression. "
The category 8.4. "Endogenic disturbances " (breakage of prerequisites 2.1.1.P.2., 2.1.1.P.1., A..2.1.1.P.1., and 2.5.P.2.) comprises a number of subcategories.
8.4.1. "Weakening of the main stock of dominants due to the shallow effective depth of the soil. "
8.4.2."Weakening of the main stock of dominants due to aging of them. "
These FDESPPs cause a failure of operation of CESPPs 2.1.1.1.1.3. "Nonpreference to herbivores " and 2.1.1.2.1.2. "Antibiosis to PPs " in dominants of forest ecosystems. When forest grows in favorable environmental conditions, the weakening takes place in the age of overmaturing. On the other hand, if the effect of 8.4.1. is potent, it takes place the phenomenon of the aged crash of forest, when dominants die in the age much less than that of normal longevity. The destructive effect gets much more, if it is accompanied by other factors of disturbance of ESPPs. Such a cooperation includes numerous subcategories of 8.2. and 8.3. Then, mortality of dominants occurs even in the young age. This is a phenomenon, which is called "the forest decline. "
The following subcategories of the category 8.4. "Endogenic disturbance " concern microevolutionary processes in PPs populations, which increase aggressiveness of them:
8.4.3. "Saltations in certain populations of phytopathogens (increase of virulence) or herbivores (increase of vitality)
8.4.4. "Microevolutionary process in herbivore populations in the conditions with weak operation of the CESPPs of the Extrinsic and Intrinsic classes.
FDESPPs 8.4.3. and 8.4.4. concern the specific threaten for ESPPs on the part of PPs. Being live organisms, they are able to evolve their capacity to counteract CESPPs. PPs are subjected by selective pressure on the part CESPPs suppressing of them. It is known the cases, when the pressure causes an appearing of taxa, which are able to counteract the factors of selection, so that FESPPs become ineffective, whereas PPs become destructive. At impact of FDESPPs 8.4.4., due to well-expressed Tolerance of foodstuff of herbivores, some ecosystems are able continually stay on the level ESPPs 3.3. "Late control." At operation of Supertolerance to herbivores, FDESPPs 8.4.4. results in a succession, i.e. composition of dominants changes.
Operation of FDESPPs 8.4.5. is known only in the ecosystems with dominance of the larch growing in the Forest-Steppe biome in south East Siberia in outbreaks of Dendrolimus sibiricus.. Here, the only significant CESPPs are .Tolerance 2.1.1.3.1.2., 2.5.6.1. and 3.5.4.1.
8.5. and 8.6. tend to wipe out CESPPs as a whole exterminating both FESPPs and prerequisites. They are so potent that ESPPs is too weak to counteract them. They are able to destroy ESPPs completely. Temporary suppressive measures are able to decrease negative effects of them. Comprehensive knowledge of FDESPPs 8.5 and 8.6.gives a possibility to preclude to some extent an operation of these impacts, and to found ways of restoring of ESPPs.
In many cases, FDESPPs operate jointly in diverse combinations that make significant difficulties in search for causes of loss of ESPPs in a concrete ecosystem. Characteristics of complexes of FDESPPs will be given in the Section 8(1).
As to plant resistance to PPs, it is used the term "durability. " This term implies a capacity of a taxon to retain the trait of resistance in wild plants over unlimited time in biocenoses, while in articenoses, the durability should be so long to compensate the expenditures for cultural practices and breeding of resistant cultivars.
The term "durability of host-plant resistance to PPs " concerns the main stock of dominants. Within this stock, a part of plants annually losses their resistance to PPs due to competition and senescence These plants are colonized by PPs, and die that, however, does not change a character of ecosystems.
There exist ecosystems with advanced Tolerance in dominants, whose ESPPs to some herbivores stays continually on the level ESPPs 3.3. "Late control. "
The disturbance of ESPPs is not immanent for ecosystems. Achievements of science allow in many situations to provide both biocenoses and articenoses by nearly all the CESPPs, which need for maintenance of their ESPPs on the level 3.1. "Proper control ". The further progress in ecosystem management will spread maintenance of ESPPs on the wider range of situations.
Ways to establish and maintain ESPPs are proceeded by people. They have been classified as CESPPs of the Anthropic class - 2.6. "Human control measures. "
Table 8. The classification of factors disturbing ecosystem stability to plant pests (FDESPPs) and their effects concerning invertebrate herbivores and phytopathogens
| Categories of FDESPPs of the first order and prerequisites of CESPPs broken due to their action | Categories of FDESPPs of the second order | CESPPs failed due to action of FDESPPs | Effects of FDESPPs on dominants | FDESPPs- possible cooperators |
| 1 | 2 | 3 | 4 | 5 |
8.1.Anthropic-1. They break prerequisites A.2.1.1.P.1., A.2.1.2.P.2.., 2.2.1.P.2., 2.2.1.P.9., 2.3.P.2., and 2.4.P.1.. due to human activity |
8.1.1. Growing of cultivars in the range of exotic herbivores r strategists |
8.1.1.1. |
8.1.1.1.1. Repeated arising of aggressive biotypes inducing outbreaks of the herbivores |
8.6.1. |
8.1.2. Growing of a single cultivar on vast areas |
8.1.2.1. |
8.1.2.1.1. Arising of high virulent pathotypes in resident phytopathogens r strategists inducing epiphytoties |
8.1.3. | |
8.1.3. Growing of cultivars in the conditions of a lack of operation of CESPPs A.2.1.2.4.1. |
8.1.3.1. |
8.1.3.1.1. Arising of high virulent pathotypes in resident phytopathogens r − strategists inducing epiphytoties |
8.1.2. | |
8.1.4. Growing of cultivars in the conditions unfavorable for expression of CESPPs A.2.1.1.2. |
8.1.4.1 |
8.1.1.4.1. Affection by phytopatogens and herbivores |
− | |
8.1.5. Growing of cultivars in the conditions enhancing of SP of PPs |
8.1.5.1. |
8.1.1.5.1. Affection by phytopathogens and herbivores |
− | |
8.1.6. Growing of cultivars with spontaneous decrease of CESPPs A.2.1.1.2.2. especially at specific weather situation |
8.1.6.1. |
8.1.6.1.1. Affection by phytopathogens |
8.3.4. | |
8.2. Anthropic-2. They break prerequisites 2.1.1.P.2., A.2.1.1.P.2., 2.1.2.P.1., A.2.1.2.P.2, A.2.1.2.4.1., 2.2.1.P.3. -.2.2.1.P.8., and 2.2.2.P.1. 2.2.2.P.3. due to human activity |
8.2.1. Cultural practices disturbing physiological state of dominants and obstacles for host-plant superevasion |
8.2.1.1. |
8.2.1.1.1. Affection by phytopathogens and herbivores |
8.2.3., |
8.2.2. Changes in the soil conditions unfavorable for dominants in forest ecosystems |
8.2.2.1. |
8.2.2.1.1. Outbreaks of stem borers |
8.2.1. | |
8.2.3. Damage of dominants by people or animals under human assistance in forest ecosystems |
8.2.3.1. |
8.2.3.1.1. Affection by phytopathogens and insect herbivores |
8.2.1. | |
8.2.4. Affection by air pollution of forest and agricultural ecosystems |
8.2.4.1. |
8.2.4.1.1. Outbreaks of stem borers in forests, signs of disease affection in agricultural ecosystems |
8.2.1. | |
8.2.5. Affection of forest ecosystems by ground fires |
8.2.5.1. |
8.2.5.1.1. Outbreaks of defoliators and stem borers |
8.2.1. | |
8.2.6. Establishing of forest stands with acorns of diverse phenological forms of the oak |
8.2.6.1. |
8.2.6.1.1. Outbreaks of defoliators of the early-spring guild on the oak |
8.2.1. | |
8.2.7. Cultural practices inducing hostile environment for natural enemies or favorable microclimate for herbivores |
8.2.7.1. |
8.2.7.1.1. Outbreaks of herbivores |
8.2.1. | |
8.3. Weather disturbances. They break prerequisites 2.3.P.1., 2.3.P.2., 2.3.P.3., and 2.3.P.4. |
8.3.1 Disturbance of water balance in dominants of forest ecosystems due to low precipitation |
8.3.1.1. |
8.3.1.1.1. Outbreaks of needle-eating defoliators and stem borers |
8.2.1. |
8.3.2. Mass coincidence of larval hatching or attacks of herbivores in older stages with favorable phases of host-plant development in dominants of forest ecosystems |
8.3.2.1. |
8.3.2.1.1. Outbreaks of defoliators of the early-spring guild of oak defoliators and the species with similar traits on coniferous trees 8.2.3., 8.2.4. |
8.2.1. | |
8.3.3. Appearing of hostile environment for natural enemies at weather extremes |
8.3.3.1. |
8.3.3.1.1. Outbreaks of defoliators of diverse guilds |
8.2.1. | |
8.3.4. Increase of aggressiveness of phytopathogens and herbivores at favorable weather in the period of their activity |
8.3.4.1. |
8.3.4.1.1. Epiphytoties and increased duration of outbreaks of herbivores and growth of their density |
8.3.3. | |
8.3.5. Increase of survivorship of PPs at favorable weather situation at dormancy |
8.3.5.1. |
8.3.5.1.1. Increase of density of herbivores and affection by phytopathogens |
8.2.1. | |
8.3.6. Weakening of dominants by the potent weather phenomena (mighty winds, sun scold or snow breakage, etc.) |
8.3.6.1. |
8.3.6.1.1. Outbreaks of stem borers |
8.2.1. | |
8.4. Endogenic disturbances. They break prerequisites 2.1.1.P.2., 2.1.1.P.1., A..2.1.1.P.1., and 2.5.P.2. |
8.4.1. Weakening of the main stock of dominants due to the shallow effective depth of the soil in forest ecosystems |
8.4.1.1. |
8.4.1.1.1. Affection by phytopathogens, increase of density of defoliators and stem borers |
8.2.1. |
8.4.2. Weakening of the main stock of dominants due to aging of them in forest ecosystems |
8.4.2.1. |
8.4.2.1.1. Affection by phytopathogens, increase of density of defoliators and stem borers |
8.2.1., | |
8.4.3. Saltations in certain populations of phytopathogens (increase of virulence) or herbivores (increase of vitality) |
8.4.3.1. |
8.4.3.1.1. Epiphytoties of phytopathogens and outbreaks of herbivores with weakly expressed periodicity |
8.2.1., | |
8.2.1., | ||||
8.4. Endogenic disturbances. They break prerequisites 2.1.1.P.2., 2.1.1.P.1., A..2.1.1.P.1., and 2.5.P.2. |
8.4.4. Microevolutionary process in herbivore populations in the conditions with weak operation of the CESPPs of the Extrinsic and Intrinsic classes |
8.4.4.1. |
8.4.4.1.1. Outbreaks of herbivores with well-expressed periodicity |
8.2.1. |
8.4.5.Growing emaciation of progeny in the conditions of weak operation of CESPPs of the Extrinsic and Intrinsic classes |
8.4.5.1. |
8.4.5.1.1.Outbreaks of herbivores with well-expressed periodicity |
- | |
8.5. Force-majeure anthropic impacts. They break prerequisites of CESPPs operating against a given taxon of PPs or CESPPs as a whole in an ecosystem |
8.5.1.Affection of ecosystems by tree crown forest fires |
8.5.1.1. Failure of all the CESPPs |
8.5.1.1.1. Outbreaks of stem borers |
− |
8.5.2.Total treatment of ecosystems by insecticides of non-selective action |
8.5.2.1. |
8.5.2.1.1. Outbreaks of defoliators |
− | |
8.5.3. Thinning of forest ecosystems in warm period of year |
8.5.3.1. |
8.5.3.1. Outbreaks of defoliators |
− | |
8.6. Force-majeure natural impacts. They break prerequisites of CESPPs operating against a given taxon of PPs or CESPPs as a whole in an ecosystem |
8.6.1. Invasion of exotic taxa of phytopathogens or herbivores, which are able to overcome CESPPs 2.1.1.2. |
8.6.1.1. |
8.6.1.1.1. Epiphytoties and outbreaks of PPs consuming biomass of the category D − I |
− |
8.6.2. Invasion of exotic taxa of herbivores, which are able to overcome CESPPs 2.2.1. |
8.6.2.1. |
8.6.2.1.1. Outbreaks of herbivores consuming biomass of the category D − II |
− | |
8.6.3. Invasion of exotic taxa of herbivores, which interact with host-plant protected by CESPPs 2.1.2.3. |
8.6.3.1. |
8.6.3.1.1. Affection of fruits by carpophagous herbivores |
− | |
8.6.4. Mass invasion of the locusts, stem borers or other insects herbivores, which are able to overcome all the local CESPPs |
8.6.4.1. Failure of CESPPs operating against a given taxon of the immigrants |
8.6.4.1.1. Destruction of plant tissues admissible for feeding by the immigrated insect herbivores |
− | |
8.6.5. Affection of ecosystems by tornado, tsunami, landslide, avalanche, etc. |
8.6.5.1. Failure of all the CESPPs |
8.6.5.1.1. In woody ecosystems, it is probable an arising of outbreaks of stem borers |
− |
