implications of silvicultural pesticides on forest soil animals, insects, fungi, and other organisms...
TRANSCRIPT
This article was downloaded by: [UQ Library]On: 11 October 2014, At: 10:03Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House,37-41 Mortimer Street, London W1T 3JH, UK
Communications in Soil Science and Plant AnalysisPublication details, including instructions for authors and subscription information:http://www.tandfonline.com/loi/lcss20
Implications of silvicultural pesticides on forest soilanimals, insects, fungi, and other organisms andramifications on soil fertility and healthFelix Ponder Jr. aa USDA Forest Service , North Central Research Station , Lincoln University , 208 Foster Hall,Jefferson City, MO, 65102, U.S.A.Published online: 05 Feb 2007.
To cite this article: Felix Ponder Jr. (2002) Implications of silvicultural pesticides on forest soil animals, insects, fungi, andother organisms and ramifications on soil fertility and health, Communications in Soil Science and Plant Analysis, 33:11-12,1927-1940, DOI: 10.1081/CSS-120004833
To link to this article: http://dx.doi.org/10.1081/CSS-120004833
PLEASE SCROLL DOWN FOR ARTICLE
Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) containedin the publications on our platform. However, Taylor & Francis, our agents, and our licensors make norepresentations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of theContent. Any opinions and views expressed in this publication are the opinions and views of the authors, andare not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon andshould be independently verified with primary sources of information. Taylor and Francis shall not be liable forany losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoeveror howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use ofthe Content.
This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in anyform to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions
IMPLICATIONS OF SILVICULTURALPESTICIDES ON FOREST SOIL ANIMALS,
INSECTS, FUNGI, AND OTHERORGANISMS AND RAMIFICATIONS ON
SOIL FERTILITY AND HEALTH
Felix Ponder, Jr.*
USDA Forest Service, North Central Research Station,
208 Foster Hall, Lincoln University, Jefferson City,
MO 65102
ABSTRACT
Sufficient information is available to show that pesticides can be
properly applied and used safely in forest management. However,
much of the information on the effects of pesticides on soil
organisms is often based on methodologies involving soil
processes (i.e., toxicology studies) rather than on individual
groups of organisms. Because more than one group of soil
organisms may carry out the same function, the effects of
pesticides on a particular group can be missed. On the other hand,
while laboratory test may show pesticide effects on one group
under an array of controlled conditions, the same organism may
respond differently in a field setting. Not much information is
available on soil organisms resulting from combining various
silvicultural herbicides or pesticides. Information is presented to
show that with few exceptions, pesticides can affect soil organism
1927
Copyright q 2002 by Marcel Dekker, Inc. www.dekker.com
*E-mail: [email protected]
COMMUN. SOIL SCI. PLANT ANAL., 33(11&12), 1927–1940 (2002)
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
populations, but the effects may be short-term with little
noticeable long-term affect on soil health and quality.
INTRODUCTION
Because of the unrelenting aggression of early succession plant species,
herbicides are likely to be a part of any cost effective approach to regenerating
old-fields to the production of fine quality hardwoods such as black walnut, red
oak, or white ash. Also, it is likely that pesticide usage is part of the history of
most old fields, those being converted to tree plantations. During the process of
selecting sites to plant trees, the decision on site quality is frequently based on
numerous physical and chemical factors. These factors may or may not be
manipulated to enhance soil quality and health; thus affecting the vigor and
health of the tree stand. Seldom, if ever, is it realized that pesticide applications to
control weeds or other pests could affect the site’s soil health and quality either
directly or through impacts on soil litter or its inhabitants. The effect of
silvicultural herbicides on soil organisms is poorly understood (1) and a
considerable amount of the data used to assess impacts are from laboratory work
rather than field trials.
The definition of soil quality and health is linked to the soil’s ability to
perform vital functions including to 1) sustain biological productivity, activity,
and diversity, 2) store and cycle nutrients and other materials, 3) partition water,
energy and solute flow, 4) filter, buffer, immobilize, and detoxify organic and
inorganic materials, and 5) support structures and protect archeological treasures
(2). Soil quality generally can be judged by soil properties that regulate their
ability to store and cycle nutrients and carbon, store and release water, and
provide favorable gas and heat exchange for specific functions. Soil
microorganisms play some role in all these soil processes, but they are
particularly important for nutrient and carbon cycling, energy flow, and nitrogen
fixation. Mycorrhizae are specialized soil fungi that penetrate secondary roots
during periods of root growth with structures called hyphae, thus becoming
extensions to roots to facilitate water and nutrient movement from soil to plant
(3–5). Alterations of physical and chemical conditions can alter the incidence of
soil borne microbes and diseases, which in turn, may affect the health of the
ecosystem and its inhabitants.
For example, habitat for small mammals that are important in distributing
fungal spores of several belowground mycorrhizal fungi (6) can be lost. Thus,
numbers, diversity, and activity of beneficial soil organisms can be reduced by
repeated removal of organic matter from a site. An examination of small mammal
digestive tracts revealed mycorrhizal fruiting bodies among the contents. Small
mammals included screws, mountain cottontails, chipmunks, ground squirrels,
PONDER1928
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
pocket gophers, mice, and voles. Ponder (7) concluded that rabbits and
grasshoppers are vectors of mycorrhizal fungi.
Further, the destruction of soil organic matter and associated degradation of
soil structure may lead to soil compaction and increased mechanical resistance to
root development (8). Another main consequence of soil organic matter loss is
the mineralization of carbon and nitrogen. Losses of carbon and nitrogen can be
substantial, as noted by Tiessen and Stewart (9), who observed net losses of 34%
carbon and 29% nitrogen over 60 years in a small grain-fallow rotation. When
cultivated soil was returned to zero tillage (no tillage) bulk density of soil in the 0
to 20 cm depth of the Ap horizon increased without substantial reduction in root
growth (10). It was suggested that roots utilized the biopores (earthworm burrows
and decayed root channels) of no tilled soil; and, hence, root growth was not
reduced. It is conceivable that young tree roots also initially utilize existing
openings in the soil during early years of growth during the conversion of
cropland and old-fields to forestland. Subsequently, during the conversion
process, soil organic matter increases. Further, soils lacking organic matter, as
well as numerous trace minerals and carbon, have very poor moisture retention.
Undoubtedly, loss of organic matter influences both soil quality and health.
Soils vary widely in their quality and capacity to perform vital functions.
Management activities affect long-term productivity and sustainability of soil by
altering soil conditions and processes. Where soil health is maintained or
improved, productivity will be maintained or improved. When soil functions are
imbalanced and soil health is degraded, plant vigor and productivity is reduced
(11). For example, long-term cultivation of croplands resulted in a reduction of
total soil organic matter (12). The purpose of this report is to discuss the
importance of soil organisms and how silvicultural management using pesticides
may affect them.
Role of Herbicides in Plantation Management
Environmentally safe, selective herbicide treatments include tree injection,
cut-stump sprays or wipes, basal sprays or wipes, directed foliar sprays, and soil-
spot and strip sprays. These treatments have the potential to control or suppress
the full range of unwanted plants when appropriate individual herbicides or tank
mixtures are used (13). Selective control can also be achieved using broadcast
applications of selective herbicides with aerial and ground systems. Changing
application rate, timing, additives, and herbicide formulation can often enhance
selectivity.
Among pesticides, chemical herbicides used in modern forestry are
vigorously tested and must meet strict standards of environmental safety and
human health protection before they are registered for use. The newer forestry
IMPLICATIONS OF SILVICULTURAL PESTICIDES 1929
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
herbicides act on biochemical processes such as photosynthesis, amino acid
pathways, and growth regulation (13). Most forestry herbicides used today
dissipate quickly by photodegradation, soil microbial biodegradation, and plant
enzyme degradation (14).
Importance of Soil Microorganisms
Soil microorganisms are increasingly the subject for study for their effect
on soil fertility and soil health, especially in the conversion of croplands into
forestlands. Soil organisms play a critical role in soil processes that regulate the
availability of nutrients and moisture in the soil for plant uptake. Their diversity
and abundance are affected by soil nutrient status, moisture, temperature, pH,
organic matter content, litter inputs, and vegetation type (15). Some soil
microorganisms have been reported to produce antibiotics or other metabolic
products, which inhibit various root pathogens (16). These organisms, their
genes, enzymes or metabolic by-products have the potential to benefit society in
the mitigation of polluted soil and for antibiotic production (17).
Another significant role of soil microorganisms is their ability to detoxify,
buffer or immobilize compounds in the soil. As integral components of the soil
foodweb, soil organisms are important food sources for many organisms at
different trophic levels.
REASONS FOR CONCERN ABOUT PESTICIDE EFFECTS ONSOIL ORGANISMS
The soil microbial community and larger inhabitants in the rhizosphere
(whole soil mass occupied by roots) are inevitably impacted on disturbed sites.
Compared to forested sites, the soil flora and fauna components beneath old-
cultivated fields are very likely to be different in both diversity and numbers.
Earthworm populations in no-till farming systems tended to be significantly
higher compared to till farming systems (18). It is important during the time of
old-field conversion to forest that management activities do not hinder the
development of these changing ecosystems. Although there is no “magic bullet”
for enhancing the restoration, there is great opportunity to use soil organisms as
“tools” (19). Also, research is now underway to identify organisms best adapted
to specific environmental and biotic conditions and to assess the potential for
managing these soil organisms. By understanding soil organisms and putting
them to wise use, land managers can better not only maintain populations of
beneficial organisms on site but also achieve greater likelihood of plantation
success. While presumed environmentally safe, selective herbicide treatments
PONDER1930
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
can be adapted to manage aboveground habitats and direct plant species
succession towards desired tree stands; there is concern that these chemicals may
affect the soil inhabitants and their functions.
Nutrient Cycling
Fine roots of developing vegetation move through the soil exuding
carbohydrates, amino acids, and other compounds that stimulate the growth of
microflora such as bacteria, antinomycetes, and fungi, which in turn produce their
own compounds that either stimulate or repel other soil organisms. Microflora are
the prime food for “grazer” herbivores such as mites, nematodes, and springtails,
which themselves fall prey to carnivores such as centipedes and spiders.
Saprophytic organisms feed on the dead remains of microbes and roots
accumulating in the rhizosphere that results in the decomposition of complex
organic molecules into basic components. Nutrients released through the
decomposition of organic matter, as well as water, may be captured and
transported to host plants by mycorrhizae (20). Mycorrhizae enhance water and
nutrient uptake not only by increasing the absorbing surface area of roots but also
through active physiological mechanisms. Ectomycorrhizal fungi (colonizes
primarily the root surface with hyphae extending into the soil) are associated with
woody species including oaks and conifers, and endomycorrhizae (colonizes the
inside of root cells with hyphae extending into the soil), which are commonly on
herbaceous species and a few woody species such as black walnut. Both groups
release enzymes that increase the availability of phosphorus to higher plants and
subsequently, its uptake (21–23). Mycorrhizae have been shown to increase
aggregation of soil particles (24).
Few nutrients leach from the soil when populations of soil organisms are
healthy and active (19). This is particularly significant for soluble forms of N
such as nitrate, which is highly soluble and mobile in the soil. As N and other
nutrients become available in the soil through decomposition of organic mater or
symbiotic and asymbiotic fixation, various soil organisms form an intricate
association to capture and assimilate N and other products into complex organic
compounds, and then slowly release them back into the soil.
Earthworms mix organic debris and applied organic and inorganic
materials, including fertilizers, lime, and pesticides, into the soil. These worms
are very important in cycling organic matter and nutrients in forest, pasture, and
no-till ecosystems (25). Although they ingest large quantities of soil and organic
matter, they excrete most of the carbon ingested (26). They form burrows that can
improve water infiltration, drainage, and aeration of the soil profile, as well as
increasing the extent and intensity of rooting. Earthworm casts and mucus
secretions tend to stabilize burrows while providing a nutrient-rich lining of the
IMPLICATIONS OF SILVICULTURAL PESTICIDES 1931
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
burrow that may further increase root proliferation. Incorporation of surface
residues by earthworms also speeds decomposition of the residues by other soil
organisms, increasing the amount and quality of the food source available,
including plant residues, can increase earthworm populations.
Nitrogen Fixation
Inadequate soil N is common on many sites. The most practical and
beneficial means of replenishing N is through biological fixation in the soil (27).
Symbiotic N fixation can add large sustained amounts of N to the soil (28).
Nitrogen-fixing plants form a mutually beneficial relationship with certain
bacteria and antinomycetes that allow for the conversion of atmospheric N into
ammonium N. The fixed N is absorbed and translocated by roots of host plants,
increasing the N concentration in living tissue, or released into the soil (29).
“Free-living” soil organisms can also contribute to the soil N supply by
asymbiotic N fixation. These organisms inhabit the soil and rotting organic
matter, and may be found in association with other soil organisms (30).
MEASURING PESTICIDE IMPACT ON SOIL ORGANISMS
Despite the extensive research on the fate of herbicides and other pesticides
in the environment, relatively little is known about how they are affected by
specific land management practices, especially over time. Many studies have
found reductions in numbers and types of soil organisms immediately following
or several years after soil disturbance, including timber harvesting, site
preparation, or prescribed fire (31–34). While short-term results are important,
managers need to know: 1) how different are these management-produced results
from results produced from natural changes such as wildfires and blowdowns and
2) do these changes affect soil productivity or sustainability. The ability or
“redundancy” of many different soil organisms to carry out soil functions and
processes makes it difficult to predict the long-term outcome. For example,
arthropod populations in a mixed-aspen forest one and two years after a wildfire
were much different than in an adjacent clearcut, but these differences were not
evident after 30 years (35).
Pesticides that are used repeatedly on a site and are slow to degrade may
result in an accumulation of pesticide residues. Some applied pesticides undergo
rapid degradation with the expectation that the ecosystem will recover
completely. While the soil seems to possess a high degree of resiliency to
management practices involving physical soil disturbances, more far reaching
effects on soil organisms may be from chemicals such as pesticides. In general,
PONDER1932
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
pesticide chemicals are not specific poisons for the particular pest they are used to
control, but are also toxic in varying degrees to other animals and man.
Crustaceans are among the most sensitive organisms. Fish are generally very
sensitive to pesticides, followed by, in order of declining sensitivity, reptiles,
birds, and mammals.
Also important in evaluating the impact of herbicides and pesticides, is their
persistence. Thirty years after the spraying DDT (dichloro diphenyl trichloroethane)
had ceased, levels of the chemical in shrews were significantly higher for sprayed
areas than for control areas (36). However, DDT levels have declined over time.
PESTICIDE DAMAGE
Of the approximately 1500 million kg of pesticides applied in the United
States, often less than 0.1% of those applied to crops reaches target pests (37).
Based on estimates by Pimentel and Levitan (37), over 99% of applied pesticides
moves into the ecosystem to be degraded by photolysis, hydrolysis, and microbial
action. Soil biota may obtain pesticides by coming in contact with them in the soil
or in plant roots (38). Once in the soil, pesticides can be adsorbed onto soil
particles, chemically bonded to other compounds in the soil, volatilized from the
soil surface into the atmosphere, move through the soil by molecular diffusion,
leached into or transported as run off in water, taken up by plant roots, or ingested
by soil fauna, thereby either entering the food chain or degraded. Partial
degradation of some pesticides can result in toxic metabolites. Some of these
chemicals may remain in the environment for a long period. Studies of pesticide
effects on soil fauna have reported increased numbers of Collembola, due
primarily to the reduction of predaceous mites; a natural enemy of Collembola
(38). These authors also conclude 1) that herbicides can greatly influence soil
invertebrates by their effects on vegetation which provides habitat and food for
many species, 2) the more persistent residues in soil do not always have as great
an influence on numbers of animals than do transient ones, and 3) most effects of
herbicides on mites and Collembola are indirect due to effects on the flora, and
generally have little effect on numbers. Many invertebrates take up pesticides
from soil into their body tissues and some, especially earthworms and mollusks,
can concentrate pesticides from soil and debris so that their tissues contain
concentrations several times greater than those in the soil in which they live (39).
When vertebrates feed on these contaminated invertebrates, they may in turn
accumulate a dose that may kill them or affect their normal activities.
Therefore, arthropods, parasites, and predators, as well as predaceous
amphibians, reptiles, birds, and mammals, may obtain and accumulate significant
quantities of pesticides by feeding on soil biota. In one investigation, for
example, the soil contained 10 ppm of DDT, earthworms contained 141 ppm, and
IMPLICATIONS OF SILVICULTURAL PESTICIDES 1933
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
the robins that were feeding on the earthworms contained 444 ppm in their brain
tissue (40). A similar situation is believed to have occurred in a midwestern city
when the robin population declined after feeding on DDT-resistant earthworms,
which had fed on fallen leaves from elm trees treated with the chemical. There is
little evidence that pesticides with relatively short half-lives and degrades
relatively quickly accumulate in tissues of invertebrates (38).
According to Lee (41), many insecticides and fungicides are toxic to
earthworms, but many herbicides are relatively harmless. All insecticides,
nematicides, mollusicides, herbicides and fungicides, once in the soil, can
potentially change populations of soil biota either directly or indirectly. Sutton
and Sheppard (42) reported that fungicide reduced mycorrhiza and subsequently
reduced aggregation of soil particles. Herbicides may greatly influence soil
invertebrate populations indirectly by their effect on vegetation, which provides
habitat and food for many of these animals.
Aside from just being able to accumulate large amounts of pesticides in
their body tissues, arthropods, plant pathogens, nematodes, and rodents have
developed resistance to pesticides. From the early 1900’s to 1980’s, a significant
number of arthropods species has developed resistance to one or more insecticide
or acarnicide (4). Georghiou and Mellon (44) reported that the number of
reported resistant species more than double in 10 years (Table 1). About 10
species of small mammals and plant attacking nematodes are known to be
resistant. Some spider mites have overcome the toxic effects of virtually every
pesticide to which they have been exposed (44).
Table 1. Increase in Number of Arthropod Species
Resistant to Pesticides
Number of Resistant Speciesa
Pesticides 1970 1980
DDT 98 229
Cyclodiens 140 269
Organophosphorus 54 200
Carbamates 3 51
Pyrethroids 3 22
Fumigants 3 7
Others 12 41
Total 313 829
a Species resistance in some part of its geographic range
does not imply that resistance is universal for any given
species [adapted from Ref. (44)].
PONDER1934
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
Table 2. Toxicities of Forest Herbicides and Other Products for Comparison; Small
Amounts for Acute Oral LD50’s Indicate a Higher Toxicity
Trade Name
Approximate
Acute Oral LD50a
(mg/kg)
Toxicity
CategorybSignal
Word
Other products
Gasoline 150 II —
Caffine 200 II —
Aspirin 1,240 III —
Baking soda 3,500 III —
Table salt 3,000 III —
Herbicides
Aatrex 4L 1,886 III Caution
Aatrex Nine-O 1,600 III Caution
Accord 5,400 IV Caution
Acme Brush Killer 2,000 III Caution
Arsenal AC .5,000 IV Caution
Banvel CST 5,000 IV Caution
Banvel 720 1,707 III Caution
Banvel 2,629 III Caution
Chopper RTU .5,000 III Caution
Escort .5,000 III Caution
Garlon 4 2,460 III Caution
Garlon 3A 2,830 III Dangerc
Krenite 24,000 IV Caution
Krenite S .5,000 IV Warningc
Oust .5,000 IV Caution
Pathway 8,000 IV Warningc
Pronone 10G .5,000 IV Caution
Rordon 5,000–6,000 IV Caution
Tordon 101 mixture 3,000 III Caution
Velpar L 7,080 IV Dangerc
Weedone CB 2,140 III Warningc
Weedone 170 2,000 III Caution
Weedone 2,4-DP 2,200 III Caution
From Ref. (13).a Unless otherwise indicated, values are for the formulated product (as in the container
before mixing).b Estimated oral amount needed to kill average person by category (II ¼ teaspoonful to an
ounce, III ¼ ounce to a pint, IV ¼ .pint).c Severe eye irritant, which increases the severity of the signal word.
IMPLICATIONS OF SILVICULTURAL PESTICIDES 1935
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
Repeated applications of the same or closely related chemicals to soils of
low adsorption have led to the development of microbial populations with an
enhanced ability to degrade the compound so that the efficacy of the pesticide is
reduced or lost (45).
Although simple acute toxicity assessments (Table 2) are possible for many
soil microorganisms, they are of dubious relevance in evaluating field responses
(46,47). In addition to mortality, changes in behavior, growth, morphogenesis,
feeding, reproduction, and even longevity must be considered. Whether laboratory
observations of these functions give valid answers are cause for concern and even
if they do, extrapolation to field predictions is filled with difficulty. Direct field
studies introduce methodological problems and are confounded by numerous
interactions with other organisms. Thus, it is perhaps best to consider pesticide
effects on soil organisms to be qualitative, rather than quantitative. Current
thinking suggests that the measurement of processes such as nitrogen
transformation and mineralization is more useful. This introduces problems
associated with naturally occurring stresses, such as drought and water logging,
which commonly reduces these activities by over 50%, before ultimate recovery.
It has therefore been suggested that assuming a microbial population would occur
in 30 days (i.e., 3 doubling times) under favorable conditions, so effects lasting less
than 30 days can be regarded as of “no ecological significance” (47). Furthermore,
Greaves and Malkomes (47) concluded that although soil processes are very
different from orthodox toxicological concepts, it is proving to be a useful
approach to the assessment of herbicide effects on microorganisms. Also, because
of rapid reproduction or the capacity of most species of soil organisms to
regenerate, initial mortality often has little permanent effect. However, initial
mortality may not always be the best estimate to measure effects on soil biota.
CONCLUSIONS
The objective explored here was to assess the importance of soil organisms
in the soil and litter and how pesticides may affect them. Only a few references
could be found in the literature on the direct effects of pesticides on microbial
activity in the soil. More cases of secondary or indirect effects were noted. The
soil microbial population is very resilient because various soil organisms carry
out similar biological processes: thus, often reducing the possibility of detecting a
change in microbial activity. In addition to using earthworms to evaluate soil
tillage effects, earthworms, can be an important tool to evaluate soil pollution and
pesticide activity. The various soil properties and different aspects of soil
management influence pesticide activity, especially leaching, which is affected
by both adsorption and rate of degradation. If the newer forestry herbicides are
applied properly, there should be little risk of damage to the ecosystem.
PONDER1936
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
REFERENCES
1. Brown, D.H. Impact of Agriculture on Bryophytes and Lichens. In
Bryophytes and Lichens in a Changing Environment; Bates, J.W., Farmer,
A.M., Eds.; Clarendon Press: Oxford, UK, 1992; 259–279.
2. Meurisse, R.T. Soil Quality and Health: Some Applications to Ecosystem
Health and Sustainability. In Proceedings of the Pacific Northwest Forest
and Rangeland Soil Organism Symposium, March 17–19, Corvalis, OR;
Meirisse, R.T., Ypsilantis, W.G., Seybold, C., Eds.; Gen. Tech. Rep. PNW-
GTR-461 U.S. Department of Agriculture, Forest Service, Pacific
Northwest Research Station: Portland, OR, 1999; 21–32.
3. Richards, B.N. The Microbiology of Terrestrial Ecosystems; John Wiley
and Sons, Inc.: New York, 1987; 399.
4. Allen, M.F. The Ecology of Mycorrhizae; Cambridge University Press:
Cambridge, 1991.
5. Harvey, A.E.; McDonald, G.I.; Jurgensen, M.F.; Larsen, M.J. Microbes:
Drivers of Long-Term Ecological Processes in Fire-Influenced Cedar–
Hemlock–White Pine Forests in the Inland Northwest. In Proceedings of
the Interior Cedar–Hemlock–White Pine Forests: Ecology and Manage-
ment, March 2–4, 1993, Spokane, WA; Baumgartner, D.A., Lotan, J.E.,
Tonn, J.R., Eds.; Washington State University: Pullman, WA, 1994;
157–163.
6. Maser, C.; Trappe, J.M.; Nussbaum, R.A. Fungal–Small Mammal
Interrelationships with Emphasis on Oregon Coniferous Forests. Ecology
1978, 59, 799–809.
7. Ponder, F., Jr. Rabbits and Grasshoppers: Vectors of Endomycorrhizal
Fungi on New Coal Mine Spoil; Res. Not. NC-250, U.S. Forest Service,
North Central Forest Experiment Station: St. Paul, MN, 1980.
8. Hamblin, A.P. The Influence of Soil Structure on Water Movement, Crop
Root Growth, and Water Uptake. Adv. Agron. 1985, 38, 95–158.
9. Tiessen, H.; Stewart, J.W.B. Particle-Size Fraction and Their Use in Studies
of Organic Matter. II. Cultivation Effects on Organic Matter Composition
in Size Fractions. Soil Sci. Soc. Am. J. 1983, 47, 509–514.
10. Ehlers, W. Penetrometer Soil Strength and Root Growth in Tilled and
Untilled Loess Soil, Proceedings of the Ninth Conference of the
International Soil Tillage Research Organization, Osijek, Yugoslavia,
1982; 458–463.
11. Waring, R.H. Imbalanced Forest Ecosystems: Assessments and Con-
sequences. For. Manag. 1985, 12, 93–112.
12. Doran, J.W.; Smith, M.S. Organic Matter Management and Utilization of
Soil and Fertilizer Nutrients. In Soil Fertility and Organic Matter as
Critical Components of Production Systems; Follett, F.F., Steward, J.W.B.,
IMPLICATIONS OF SILVICULTURAL PESTICIDES 1937
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
Cole, C.V., Eds.; SSSA Spec. Publ. No. 19, Soil Science Society of
America: Madison, WI, 1987; 53–72.
13. McMahon, C.K.; Miller, J.H.; Thomas, D.F. Role of Low Impact Pesticide
Treatments in Ecosystem Management. In The 15th Annual Forest
Vegetation Management Conference, Redding, CA, Jan 25–27, 1994;
171–172.
14. Thompson, D.G.; Staznik, B.; Fontaine, D.D.; Mackay, T.; Oliver, G.R.;
Troth, J. Fate of Triclopyr Ester (RELEASE) in a Boreal Forest Stream.
Environ. Toxicol. Chem. 1991, 10, 619–632.
15. Harvey, A.E.; Larsen, M.J.; Jurgensen, M.F. Clearcut Harvesting and
Ectomycorrhizae: Survival of Activity on Residual Roots and Influence on
a Bordering Forest Stand in Western Montana. Can. J. For. Res. 1980, 10,
300–303.
16. Azcon-Aguilar, C.; Barea, J.M. Interactions Between Mycorrhizal Fungi
and Other Rhizosphere Microorganisms. In Mycorrhizal Functioning: An
Integrative Plant–Fungal Process; Allen, M.F., Ed.; Chapman & Hall:
New York, 1992; 163–198.
17. Jurgensen, M.F. Soil Organisms: Functions and Processes—Management
Implications: A Synopsis. In Proceedings of the Pacific Northwest Forest
and Rangeland Soil Organism Symposium, March 17–19, Corvalis, OR;
Meirisse, R.T., Ypsilantis, W.G., Seybold, C., Eds.; Gen. Tech. Rep. PNW-
GTR-461, U.S. Department of Agriculture, Forest Service, Pacific
Northwest Research Station: Portland, OR, 1999; 205–210.
18. Jordan, D.; Stecker, J.A.; Cacnio-Hubbard, V.N.; Li, F.; Gantzer, C.J.;
Brown, J.R. Earthworm Activity in No-Tillage and Conventional Tillage
Systems in Missouri Soils: A Preliminary Study. Soil Biol. Biochem. 1997,
29, 489–491.
19. Amaranthus, M.P.; Cazares, E.; Perry, D.A. The Role of Soil Organisms in
Restoration. In Proceedings of Pacific Northwest Forest and Rangeland
Soil Organism Symposium, March 17–19, Corvalis, OR; Meirisse, R.T.,
Ypsilantis, W.G., Seybold, C., Eds.; Gen. Tech. Rep. PNW-GTR-461, U.S.
Department of Agriculture, Forest Service, Pacific Northwest Research
Station: Portland, OR, 1999; 179–189.
20. Harley, J.L.; Smith, S.E. Mycorrhizal Symbioses; Academic Press: London,
1983.
21. Alexander, I.J.; Hardy, K. Surface Phosphatase Activity of Sitka Spruce
Mycorrhizas from a Serpentine Site. Soil Biol. Biochem. 1981, 13,
301–305.
22. Moorman, T.; Reeves, F.B. The role of Endomycorrhizae in Revegetation
Practices in the Semi-Arid West. II. A Bioassay to Determine the Effect of
Land Disturbance on Endomycorrhizal Populations. Am. J. Bot. 1979, 66,
14–18.
PONDER1938
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
23. Ponder, F., Jr. Growth and Mycorrhizal Development of Potted White Ash
and Black Walnut Fertilized by Two Methods. Can. J. Bot. 1984, 62,
509–512.
24. Koske, R.E.; Sutton, J.C.; Sheppard, B.R. Ecology of Endogone in Lake
Huron Sand Dunes. Can. J. Bot. 1975, 53, 87–93.
25. Kladivko, E.J.; Timmenga, H.J. Earthworms and Agricultural Manage-
ment. In Rhizosphere Dynamics; Box, J.E., Hammond, L.C., Eds.;
Westview Press, Inc.: Oxford, 1990; 192–216.
26. Bolton, P.J.; Phillipson, J. Burrowing, Feeding, Egestation and Energy
Budgets of Allolobophora rosea (Savigny) (Lumbricidae). Oecologia 1976,
23, 225–245.
27. Gordon, J.C.; Avery, M.E. Improving Tree Crops Using Microorganisms in
Designed Systems. In Attributes of Tree Crop Plants; Cannell, M.G.R.,
Jackson, J.E., Eds.; Institute of Terrestrial Ecology, Abbots Ripton: Hunts,
UK, 1985; 316–326.
28. Erdman, L.W. Legume Inoculation; USDA Farmers Bull. No. 2003,
Government Printing Office: Washington, DC, 1967.
29. Tarrant, R.F.; Trappe, J.M. The Role of Alder in Improving the Forest
Environment. Plant Soil 1971, Special Volume, 335–348.
30. Amaranthus, M.P.; Molina, R.; Perry, D.A. Soil Organisms, Root Growth
and Forest Regeneration. Forestry on the Frontier Proceedings; National
Society of American Foresters National Convention: Spokane, WA, 1990;
89–93.
31. Bissett, J.; Parkinson, D. Long-Term Effects of Fire on the Composition and
Activity of the Soil Microflora of a Subalpine, Coniferous Forest. Can.
J. Bot. 1980, 58, 1704–1721.
32. Blair, J.M.; Crossley, D.A. Litter Decomposition, Nitrogen Dynamics and
Litter Microarthropods in a Southern Applachain Hardwood Forest 8 Years
Following Clearcutting. J. Appl. Ecol. 1988, 25, 683–698.
33. Pietikainen, J.; Fritze, H. Clear-Cutting and Prescribed Burning in
Coniferous Forest: Comparison of Effects on Soil Fungal and Total
Microbial Biomass, Respiration Activity and Nitrification. Soil Biol.
Biochem. 1995, 27, 101–109.
34. Prescott, C.E. Effects of Clearcutting and Alternative Silvicultural Systems
on Rates of Decomposition and Nitrogen Mineralization in a Coastal
Montane Coniferous Forest. For. Ecol. Manag. 1997, 95, 253–260.
35. Spence, J.R.; Buddle, C.M.; Gandhi, K.J.K.; Langor, D.W.; Volney,
W.J.A.; Hammond, H.E.J.; Pohl, G.R. Invertebrate Biodiversity, Forestry
and Emulation of Natural Disturbance: A Down to Earth Perspective. In
Proceedings of the Pacific Northwest Forest and Rangeland Soil Organism
Symposium, March 17–19, Corvalis, OR; Meirisse, R.T., Ypsilantis, W.G.,
Seybold, C., Eds.; Gen. Tech. Rep. PNW-GTR-461, U.S. Department of
IMPLICATIONS OF SILVICULTURAL PESTICIDES 1939
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014
Agriculture, Forest Service, Pacific Northwest Research Station: Portland,
OR, 1999; 80–90.
36. Perkins, D.W.; Hodgman, T.P.; Owen, R.B.; Dimond, J.B. Long-Term
Persistence of DDT in Shrews, Soricidae, from Maine. Can. Field-Nat.
1998, 112, 393–399.
37. Pimentel, D.; Levitan, L. Pesticides: Amounts Applied and Amounts
Reaching Pests. BioScience 1986, 36, 86–91.
38. Edwards, C.A.; Thompson, A.R. Pesticides and the Soil Fauna. Residue
Rev. 1973, 45, 1–79.
39. Headley, J.C.; Lewis, J.N. The Pesticide Problem: An Economic Approach
to Public Policy; The Johns Hopkins Press: Baltimore, MD, 1967; 141.
40. Hunt, L.B. Kinetics of Pesticide Poisoning in Dutch-Elm Disease Control.
I.S. Fish Wildl. Serv. Circ. 1965, 226, 12–13.
41. Lee, K.E. Earthworms: Their Ecology and Relationship with the Soil and
Land Use; Academic Press: New York, 1985.
42. Sutton, J.C.; Sheppard, B.R. Aggregation of Sand-Dune Soil by
Endomycorrhizal Fungi. Can. J. Bot. 1976, 54, 326–333.
43. Dover, M.J.; Craft, B.A. Pesticide Resistance and Public Policy.
BioScience 1986, 36, 78–85.
44. Georghiou, G.P.; Mellon, R.B. Pesticide Resistance in Time and Space. In
Pest Resistance to Pesticides; Georghiou, G.P., Saito, T., Eds.; Plenum
Press: New York, 1983; 1–46.
45. Kaufman, D.D.; Katan, Y.; Edwards, D.F.; Jordan, E.G. Microbial
Adaptation and Metabolism of Pesticides. In Agriculture Chemicals of the
Future; Hilton, J.L., Ed.; BARC Symp. No. 8, Rowan and Allanhold:
Totowa, NJ, 1983; 35.
46. Domsch, K.H.; Jagnow, G.; Anderson, T.H. An Ecological Concept for the
Assessment of Side Effects of Agrochemicals on Soil Microorganisms.
Residue Rev. 1983, 86, 65–105.
47. Greaves, M.P.; Malkomes, H.P. Effects on Soil Microflora. In Interactions
Between Herbicides and the Soil; Hance, B.J., Ed.; Academic Press:
London, 1980; 48.
PONDER1940
Dow
nloa
ded
by [
UQ
Lib
rary
] at
10:
03 1
1 O
ctob
er 2
014