Download stability of terrestrial ecosystems as to pest organisms

G.I. Vasechko STABILITY OF TERRESTRIAL ECOSYSTEMS TO PLANT PESTS: AN AXIOMATIC APPROACH.
PART II. SUBSTANTIATION OF THE AXIOMS PROPOSED IN THE PART I
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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).
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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
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."
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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
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.
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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
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."
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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
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
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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
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.
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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
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
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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
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.
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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
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
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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
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."
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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
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.
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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
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.
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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
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).
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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
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