Potential impact of the leaf‐cutting ant Acromyrmex lobicornis on conifer plantations in northern Patagonia,Argentina

• 1 The economic losses associated with crop damage by invasive pests can be minimized by recognizing their potential impact before they spread into new areas or crops.
• 2 We experimentally evaluated the preferences of the leaf‐cutting ant Acromyrmex lobicornis (Hymenoptera: Formicidae) for the most common conifer species commercially planted in northern Patagonia, Argentina. The areas of potential forest interest in this region and the geographical range of this ant overlap. We performed field preference tests and monitored the level of ant herbivory on planted conifer seedlings next to nests.
• 3 Acromyrmex lobicornis preferred some conifer species and avoided foraging on others. Pseudotsuga menziesii and Austrocedrus chilensis were the less preferred species, Pinus ponderosa and Pinus contorta were the most preferred by A. lobicornis.
• 4 The item mostly selected by ants was young needles from P. contorta. This species was also the pine mostly defoliated. Seedlings without ant‐exclusion showed a mean±SE of 60±5% defoliation during the sampling period. Pinus ponderosa was less defoliated; control seedlings showed a mean±SE of 8.5±1% of leaf damage in the sampling period.
• 5 The present study shows how the use of simple field tests of leaf‐cutting ant preferences could allow an improved selection of appropriate conifer species for future plantations in areas where leaf‐cutting ants are present.

     

Economic Injury Level and Economic Threshold as required by Forest Stewardship Council for management of leaf-cutting ants in forest plantations

1. The most commonly used method to control insect pests, such as leaf-cutting ants (LCA) in forest plantations, is the application of chemical insecticides, but their use was forbidden in plantations certified by the Forest Stewardship Council because of their negative effect on the environment.
2. A 5-year derogation of their prohibition was obtained in 2016 in Argentina, on the condition that forestry companies reduce pesticide doses by 5% per year and inform the Economic Injury Level (EIL).
3. The EIL and the Economic Threshold (ET) were estimated for the first time for the Acromyrmex genus in a Salicaceae forestation. As a model, we used the damage caused by the most detrimental LCA species of Argentina (Acromyrmex lundii) in the first, most critical year of a willow plantation (Salix nigra) in the lower basin of the Paraná River.
4. The EIL (8–15 nest/ha) was useful to demonstrate the real magnitude of the damage caused by A. lundii, which justified the need for its control. The ET for profitable production (∼3 nests/ha) would not be operatively applicable in a long-term management program in our study site. This methodology could be extrapolated to other Neotropical forest plantations attacked by Acromyrmex species.

     

Tree species identity and forest composition affect the number of oak processionary moth captured in pheromone traps and the intensity of larval defoliation

1. Pure forests are often seen as being more prone to damage by specialist pest insects than mixed forests, and particularly mixed forests associating host and nonhost species. We addressed the effect of tree diversity on oak colonization and defoliation by a major specialist pest, the oak processionary moth (OPM)
2. We quantified the number of male OPM moths captured and larval defoliation in pure stands of two oak host species (Quercus robur and Quercus petraea) and in mixed stands associating the two oak species or each oak species with another nonhost broadleaved species. We conducted two complementary studies to test the effect of host species and stand composition: (i) we used pheromone trapping to compare the number of males OPM captured throughout the distribution of oak hosts in France and (ii) we noted the presence of OPM nests and estimated defoliation in mature forests of north‐eastern France.
3. Oak species and stand composition significantly influenced the number of male OPM captured and defoliation by OPM larvae. Quercus petraea was consistently more attractive to and more defoliated by OPM than Q. robur. Both oak trees were attacked more in pure stands than in mixed stands, in particular mixed stands associating oaks with another (nonhost) broadleaved species.
4. The results of the present study support the view that mixed forests are more resistant to specialist pest insects than pure stands, and also indicate that this trend depends on forest composition. Our study provides new insights into OPM ecology and has potential implications for forest management, including the management of urban forests where OPM causes serious human health issues.

     

Resource availability and growth responses to defoliation in seedlings of three early-successional,tropical, woody species

A growth experiment was conducted using seedlings of three early-successional, tropical, woody species:Dillenia suffruticosa (Dilleniaceae),Macaranga heynei (Euphorbiaceae) andTrema tomentosa (Ulmaceae). These species are characteristic of different positions along a soil fertility gradient in Singapore, withD. suffruticosa being the least andT. tomentosa the most demanding of high nutrient availability. The seedlings were grown in vermiculite at either low or high nutrient availabilities supplied by watering with different concentrations of a commercial plant food. Half the seedlings were subjected to a 50% defoliation at the start of the experiment by cutting off the distal half of each leaf. After 9 weeks the plants were harvested.Macaranga heynei andT. tomentosa seedlings showed no significant difference in parameters of growth such as total dry weight and total leaf area between the defoliated and control seedlings. The seedlings compensated completely for the loss of leaf area.Dillenia suffruticosa did show significant reductions in growth in some parameters due to defoliation, and these were more pronounced under the high nutrient treatment. These findings support the hypothesis that plants characteristic of resource-rich sites can readily recover from herbivory through fast growth, probably associated with a rapid turnover of leaves, whereas species of resource-poor habitats cannot easily replace losses due to herbivory and are adversely effected by defoliation.     

Impact of defoliation intensities on plant biomass,nutrient uptake and arbuscular mycorrhizal symbiosis in Lotus tenuis growing in a saline‐sodic soil

The impact of different defoliation intensities on the ability of Lotus tenuis plants to regrowth, mobilise nutrients and to associate with native AM fungi and Rhizobium in a saline‐sodic soil was investigated. After 70 days, plants were subjected to 0, 25, 50, 75 and 100% defoliation and shoot regrowth was assessed at the end of subsequent 35 days. Compared to non‐defoliated plants, low or moderate defoliation up to 75% did not affect shoot regrowth. However, 100% treatment affected shoot regrowth and the clipped plants were not able to compensate the growth attained by non‐defoliated plants. Root growth was more affected by defoliation than shoot growth. P and N concentrations in shoots and roots increased with increasing defoliation while Na⁺ concentration in shoots of non‐defoliated and moderately defoliated plants was similar. Non‐defoliated and moderately defoliated plants prevented increases of Na⁺ concentration in shoots through both reducing Na⁺ uptake and Na⁺ transport to shoots by accumulating Na⁺ in roots. At high defoliation, the salinity tolerance mechanism is altered and Na⁺ concentration in shoots was higher than in roots. Reduction in the photosynthetic capacity induced by defoliation neither changed the root length colonised by AM fungi nor arbuscular colonisation but decreased the vesicular colonisation. Spore density did not change, but hyphal density and Rhizobium nodules increased with defoliation. The strategy of the AM symbiont consists in investing most of the C resources to preferentially retain arbuscular colonisation as well as inoculum density in the soil.     

Impacts of defoliation by tussock moths (Orgyia vetusta) on the growth and reproduction of bush lupine (Lupinus arboreus)

Abstract.

• 1 Outbreaks of tussock moths (Orygia verusta Bdv.) on bush lupines (Lupinus arboreus Sims) may be very intense, although spatially localized, and may last >10 years. To understand better how such outbreaks may persist, we defoliated lupine bushes using variable realistic numbers of tussock moth larvae, and measured the immediate and delayed impacts of defoliation on the growth and seed production of lupine bushes.
• 2 Immediate effects of defoliation included the production of new leaves that were significantly smaller in diameter and biomass, and lower in water content, than the new leaves of undamaged bushes. Also, seed output was reduced by up to 80% on heavily damaged bushes.
• 3 In the longer term, surviving bushes recovered remarkably well, despite having received the full range of possible (survivable) insect densities. After 1 or 2 years’rapid growth, surviving bushes were not affected in terms of height, basal stem diameter or volume by their previous defoliation. However, there was a significant tendency for juvenile bushes to produce more seeds the more heavily they had been attacked the previous year.
• 4 The ability of the host plant to recover, in terms of its biomass, helps to explain the sustained nature of the insect outbreak.

     

Growth,nutrition and gas exchange of Pinus resinosa following artificial defoliation

Summary In three experiments, red pine (Pinus resinosa Ait.) seedlings and trees were subjected to artificial defoliations of varying intensities and subsequent growth, gas exchange and nutritional responses were monitored. In Experiment 1, 2-year-old seedlings received 0, 1 or 2 50% defoliations during a single growing season and were maintained in 1 of 3 low nutrient supply treatments. In Experiment 2, response of 4-year-old seedlings was monitored in the year following 0, 25, 50 or 75% defoliation, while in Experiment 3, response of 11-year-old trees was measured 1 year after being defoliated by 0, 33 or 66%. Regardless of intensity of defoliation, or plant size, clipped plants made qualitatively similar allocational and metabolic adjustments over time. First, leaf diffusive conductance and rates of net photosynthesis were stimulated, especially by light to intermediate defoliation. However, there was no effect of defoliation on foliar nitrogen concentration, and elevated gas exchange rates apparently resulted from altered root-shoot dynamics. Second, allocation of new biomass was preferentially shifted towards foliage at the expense of roots, gradually restoring (but undershooting or overshooting) the ratio of foliage: roots of control plants. During the period when foliage: root balance was being restored, the stimulation of needle gas exchange rates disappeared. Plants defoliated by 25% overcompensated in terms of whole plant growth (were larger at harvest than controls), due to shifts in allocation and enhanced photosynthesis. Defoliated plants also stored a proportionally greater share of their carbohydrate reserves in roots than did control plants, even 1 year after clipping.     

Defoliation increases carbon limitation in ectomycorrhizal symbiosis of <Emphasis Type="Italic">Betula pubescens</Emphasis>

Boreal forest trees are highly dependent on root-colonizing mycorrhizal fungi. Since the maintenance of mycorrhizal symbiosis implies a significant carbon cost for the host plant, the loss of photosynthetic leaf area due to herbivory is expected to reduce the host investment in mycorrhizae. We tested this hypothesis in a common garden experiment by exposing ectomycorrhizal white birch (Betula pubescens Ehrh.) seedlings to simulated insect defoliation of 50 or 100% intensity during either the previous or the current summer or repeatedly during both seasons before harvest. The shoot and root growth of the seedlings were distinctly reduced by both 100% defoliation and repeated 50% defoliation, and they were more strongly affected by previous-year than current-year defoliation. The root to shoot ratio significantly decreased after 100% defoliation, indicating reduced proportional allocation to the roots. Ergosterol concentration (i.e. fungal biomass) in the fine roots decreased by 100% defoliation conducted either in the year of harvest or in both years. No such decrease occurred following the 100% defoliation conducted in the previous year, indicating the importance of current photosynthates for fungal symbionts. The trend was similar in the colonization percentage of thick-mantled mycorrhizae in the roots, the most marked decline occurring in the repeatedly defoliated seedlings. The present results thus support the prediction that the plant investment in ectomycorrhizae may decline as a response to foliage loss. Moreover, the colonization percentage of thick-mantled mycorrhizae correlated positively with the ratio of leaf to heterotrophic plant biomass in the defoliated birch seedlings, but not in the control ones. This tends to indicate a stronger carbon limitation of ectomycorrhizal colonization in defoliated seedlings.     

Impact of simulated herbivory on water relations of aspen (<Emphasis Type="Italic">Populus tremuloides</Emphasis>) seedlings: the role of new tissue in the hydraulic conductivity recovery cycle

Physiological mechanisms behind plant–herbivore interactions are commonly approached as input–output systems where the role of plant physiology is viewed as a black box. Studies evaluating impacts of defoliation on plant physiology have mostly focused on changes in photosynthesis while the overall impact on plant water relations is largely unknown. Stem hydraulic conductivity (k _h), stem specific conductivity (k _s), percent loss of hydraulic conductivity (PLC), CO₂ assimilation (A) and stomatal conductance (g _s) were measured on well-irrigated 1-month-old Populus tremuloides (Michx.) defoliated and control seedlings until complete refoliation. PLC values of defoliated seedlings gradually increased during the refoliation process despite them being kept well irrigated. k _s of defoliated seedlings gradually decreased during refoliation. PLC and k _s values of control seedlings remained constant during refoliation. k _s of new stems, leaf specific conductivity and A of leaves grown from new stems in defoliated and control seedlings were not significantly different, but g _s was higher in defoliated than in control seedlings. The gradual increase of PLC and decrease of k _s values in old stems after defoliation was unexpected under well-irrigated conditions, but appeared to have little impact on new stems formed after defoliation. The gradual loss of conductivity measured during the refoliation process under well-irrigated conditions suggests that young seedlings of P. tremuloides may be more susceptible to cavitation after herbivore damage under drought conditions.     

Effect of partial defoliation during the vegetative phase on subsequent growth and grain yield of maize

Maize was grown in two densities, 2–47 or 4–94 plants m^-2, and the following treatments imposed: untreated, plants partly defoliated 51 days after sowing, and alternate plants in a row partly defoliated 44 days after sowing. Plants flowered about 82 days after sowing. Leaf area was decreased by 60–64% by defoliation on day 51. Defoliation resulted in decreases in grain yield and grain number of 6–17%, though when alternate plants were defoliated in the higher density there was a substantial decrease in yield and number of grains in defoliated plants, which was largely offset by an increase in adjacent intact plants. When plants were defoliated on day 51 subsequent growth in leaf area was similar to, and that in leaf weight nearly as large as that in untreated plants, while increase in stem weight was substantially less than in untreated plants. By the time of flowering untreated and defoliated plots differed by c. 30% in leaf area. Increments of dry matter after flowering differed by c. 15% between untreated and defoliated plots. The fraction of these increments which entered the grain was c. 90% in both untreated and defoliated plots. When alternate plants in the row were partly defoliated on day 44 their subsequent increase in leaf area was probably 5–16% less than that of the adjacent intact plants. Increments of dry matter after flowering of plots with alternate plants defoliated were 93–95 % of those of untreated plots; leaf efficiency after flowering was slightly greater than in untreated plots. The fraction of the dry matter increment after flowering which entered the grain was c. 88 % in both intact and defoliated plants of the small density, but was 94% in intact plants and 86% in defoliated plants of the large density.     

Lepidoptera‐specific insecticide used to suppress gypsy moth outbreaks may benefit non‐target forest Lepidoptera

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Defoliation of Brassica napus increases severity of blackleg caused by Leptosphaeria maculans: implications for dual-purpose cropping

Canola (Brassica napus) crops for grazing and grain (dual-purpose) production provide an economic break-crop alternative for dual-purpose cereals in Australian mixed farming systems. Infection by Leptosphaeria maculans is the most prevalent disease in Australian canola crops with airborne inoculum released throughout the autumn and winter when crops are grazed. Glasshouse and field experiments were conducted to investigate the effect of mechanical defoliation (simulated grazing) on disease severity at plant maturity. In glasshouse experiments, stem canker severity increased from 4% to 24% in severely defoliated plants, but light defoliation had no effect compared with undefoliated control plants. Disease severity was increased with defoliation in all field experiments. Defoliation increased crown canker severity from 22.6% to 39.3% at Wagga Wagga and from 3.0% to 7.1% at Canberra and lodging from 9.6% to 11.9% at Naracoorte in the same set of cultivars assessed at each site. The increase in disease severity with defoliation was less in canola lines with moderate to high levels of stem canker resistance. Plants defoliated before stem elongation tended to develop less disease than those defoliated during the reproductive phase of plant growth. These findings suggest that the impact of grazing on L. maculans infection of canola crops can be minimised by sowing cultivars with a high level of stem canker resistance and grazing during the vegetative stage of plant growth prior to stem elongation. Further research is required to determine whether these management strategies are applicable in canola crops defoliated by grazing animals.     

Analytical studies on the responses of sunflower (Helianthus annuus) to various defoliation treatments

The effects of defoliation treatments on plant growth in sunflower (Helianthus annuus) were studied in the field. Four defoliation treatments, 0 (control), 37.4, 56.1 and 93.4% of the total leaf dry weight, were applied to plants that had small third leaves. Decreased leaf weight/whole plant weight (F/W) ratios in defoliated plants rapidly recovered to almost the same ratio as that observed in the control within 12 to 16 days after defoliation according to the degree of defoliation. The mechanism involved in the recovery of the F/W ratio in defoliated plants mainly consisted of three parameters: enhancement of (1) carbon distribution ratios in the leaves, (2) photosynthetic activity in the remaining leaves, and (3) retranslocation of carbon from the stem and/or roots to leaves. Inhibitive effects of defoliation on relative growth rate and net assimilation rate were seen at an early stage, but subsequently both rates became larger in defoliated plants than in controls. Defoliated plants tended to show rapid development and expansion of new leaves, and to show increased specific leaf area and protein synthesis in individual leaves. The sugar content of leaves in defoliated plants was higher than that in controls, while the content in both stem and roots was lower. These responses seem to be advantageous for development of the photosynthetic system. Heights of defoliated plants were clearly depressed according to the degree of defoliation, and this was attributed largely to differences in the elongation rates of the internodes resulting from defoliation.     

Levels of polyphenol oxidase activity in leaves of Milicia seedlings is indicative of the resistance to Phytolyma lata-induced gall disease

1. Phytolyma lata-induced galling has remained one of the foremost challenges limiting the regeneration of Milicia species in nurseries and degraded forests in West Africa. Although planting of Milicia progenies resistant to Phytolyma attack has been identified as an opportunity for long-term control of the pest, the absence of biomarkers for early detection of resistance has limited the selection of resistant seedlings for planting.
2. We compared phenolic metabolites and oxidative enzymes in resistant and susceptible Milicia progenies with the aim of identifying potential biomarkers for resistance to Phytolyma-induced galling in seedlings.
3. Levels of total phenolics and ortho-dihydroxyphenolics did not significantly differ between the susceptible and resistant saplings or among seedlings. Similarly, peroxidase activity did not differ among sapling progenies or among seedlings. Thus, no clearly defined role could be ascribed to phenolics and peroxidase in resistance of Milicia species to P. lata-induced galling. However, polyphenol oxidase (PPO) activity was over two-fold higher in the resistant than the tolerant or susceptible progenies, suggesting that higher levels of PPO activity corresponded with an increased level of resistance to galling.
4. With further validation, PPO activity in leaves of Milicia seedlings can be used as a biomarker of resistance to Phytolyma-induced galling.

     

Interactive effects of defoliation and an AM fungus on plants and soil organisms in experimental legume–grass communities

We established a 13‐week greenhouse experiment based on replicated microcosms to test whether the effects of defoliation on grassland plants and soil organisms depend on plant species composition and the presence of arbuscular mycorrhizal (AM) fungi. The experiment constituted of three treatment factors – plant species composition, inoculation of an AM fungus and defoliation – in a fully factorial design. Plant species composition had three levels: (1) Trifolium repens monoculture (T), (2) Phleum pratense monoculture (P) and (3) mixture of T. repens and P. pratense (T+P), while the AM inoculation and the defoliation treatment had two levels: (1) no inoculation of AM fungi and (2) inoculation of the AM fungus Glomus claroideum BEG31, and (1) no trimming, and (2) trimming of all plant material to 6 cm above the soil surface three times during the experiment, respectively. At the final harvest, AM colonization rate of plant roots differed between the plant species compositions, being on average 45% in T, 33% in T+P and 4% in P. Defoliation did not affect the colonization rate in T but raised the rate from 1% to 7% in P and from 20% to 45% in T+P. Shoot production and standing shoot and root biomass were 48%, 85% and 68% lower, respectively, in defoliated than in non‐defoliated systems, while the AM fungus did not affect shoot production and root mass but reduced harvested shoot mass by 8% in non‐defoliated systems. Of the plant quality attributes, defoliation enhanced the N concentration of harvested shoot biomass by 129% and 96% in P and T+P, respectively, but had no effect in T, while the C concentration of shoot biomass was on average 2.7% lower in defoliated than in non‐defoliated systems. Moreover, defoliation reduced shoot C yield (the combined C content of defoliated and harvested shoot biomass) on average by 47% across all plant species compositions and shoot N yield by 37% in T only. In contrast to defoliation, the AM fungus did not affect shoot N and C concentrations or shoot N yield, but induced 10% lower C yield in non‐defoliated systems and 17% higher C yield in defoliated T. In roots, defoliation led to 56% and 21% higher N concentration in P and T+P, respectively, and 28% higher C concentration in P, while the mycorrhizal fungus lowered root N concentration by 9.7% in defoliated systems and had no effect on root C concentrations. In the soil, the nematode community was dominated by bacterivores and the other trophic groups were found in a few microcosms only. Bacterivores were 45% more abundant in defoliated than in non‐defoliated systems, but were not affected by plant species composition or the AM fungus. Soil inorganic N concentration was significantly increased by defoliation in T+P, while the mycorrhizal fungus reduced NH₄–N concentration by 40% in T. The results show that defoliation had widespread effects in our experimental systems, and while the effects on plant growth were invariably negative and those on bacterivorous nematodes invariably positive, most effects on plant C and N content and soil inorganic N concentration varied depending on the plant species present. In contrast, the effects of defoliation did not depend on the presence of the AM fungus, which suggests that while the relative abundance of legumes and grasses is likely to have a significant role in the response of legume–grass communities to defoliation, the role of AM fungi may be less important. In line with this, the AM fungus had only a few significant effects on plant and soil attributes in our systems and each of them was modified by defoliation and/or plant species composition. This suggests that the effects of AM fungi in legume–grass communities may largely depend on the plant species present and whether the plants are grazed or not.     

Density- and growth stage-dependent responses to defoliation in two rhizomatous grasses

Summary Responses to defoliation were studied in two tallgrass prairie perennials (Andropogon gerardii and Panicum virgatum) established from seed at three densities. P. virgatum was also grown from transplanted rhizomes of established clones. Plants of both species displayed a continuum of responses to defoliation, from large reductions in biomass, tillering and seed production to significant increases in one or more performance measures. In crowded populations, defoliation shifted plants into subordinate positions within the competitive hierarchy. Plants competing intraspecifically and those that were initially small suffered more from defoliation than either plants grown at low density or those that were larger than their neighbors. At the highest plant density, the effects of defoliation or initial plant size were overshadowed by the effects of crowding. When defoliated and grown at similar densities, P. virgatum and A. gerardii grown from seed showed large reductions in biomass, seed production, and new rhizome production, but established P. virgatum ramets grown from rhizomes showed increases in these performance measures. Thus, herbivory may be particularly detrimental to P. virgatum during juvenile stages before perennating organs have developed. Overcompensation of P. virgatum clones in response to defoliation only occurred if all ramets within the clone were defoliated. In clones containing both defoliated and undamaged ramets, there were no differences in their performance, suggesting that genets are capable of integrating the effects of differential defoliation among shoots. Defoliated P. virgatum clones allocated a smaller fraction of their total biomass to new rhizomes, indicating that the short-term regrowth response following defoliation may incur a longer-term cost associated with gradual reduction in biomass of the perennating organs and reduced genet success.     

Colony size,density and distribution of the leaf-cutting ant,Acromyrmex octospinosus (Reich) in cultivated fields

• 1 Fields of citrus, cacao and rubber in Trinidad were surveyed to measure the number and density of nests of Acromyrmex octospinosus. The greatest density (=153 nests/ha) was in an undisturbed field of young cacao; in established crops density ranged from 3–36 nests/ha.
• 2 Mounds of new nests appeared throughout the year, but most often 4–10 months after the main nuptial flights. Nests reached maximum size, with up to 17 holes, about three years after foundation.
• 3 The ratio of workers to brood was between 5 and 9: 1 for most of the year but it increased to 27: 1 as the sexuals developed. A medium-large colony produced 200–300 queens and males/year; the largest single nest contained about 14,000 individuals.
• 4 The average intake of leaf fragments into laboratory nests was 1 fragment/6.5 individuals. It increased to 1 fragment/2.1 individuals when sexuals were present. Foragers collecting from fields of established cacao preferred flowers to leaves.
• 5 In fields with crops planted on cambered beds, nests were concentrated on the dry edges of drains in the dry season, but they were moved to drier ground in the wet season. Nests were aggregated in 13 of the 16 fields examined, the remaining three fields were exceptionally well drained.
• 6 In newly planted cacao an estimated 6—17% of plants were defoliated and destroyed. In established cacao ants collected about 3000 flowers/ha each day. Laboratory data suggest that ants could defoliate about 25 three-year old citrus trees in a year.

     

Net photosynthesis,root respiration,and regrowth of Bouteloua gracilis following simulated grazing

Summary Net photosynthesis (P_N), root respiration (R_R), and regrowth of Bouteloua gracilis (H.B.K.) Lag. were examined in the laboratory over a 10-day period following clipping to a 4-cm height to simulate grazing by large herbivores. Net photosynthesis rates of tissue remaining immediately following defoliation were only about 40% as great as preclipping rates. Three days after clipping, P_N rates of defoliated plants had increased to values about 21% greater (per unit leaf area) than those of unclipped controls and remained at that level through Day 10. No statistically significant changes in R_R occurred following defoliation. Biomass of unclipped plants nearly doubled during the 10-day study period, while that of defoliated plants increased 67%. Over half the new growth of defoliated plants was allocated to new leaf blades and only 18% to new roots, while only 33% of the new growth of control plants was allocated to new leaf blades but 29% went to new roots. As a consequence of increased P_N rates and increased carbon allocation to synthesis of additional photosynthetic tissue following defoliation, net CO₂ uptake per plant increased from 9% to 80% of that of the controls from Day 0 through Day 10.     

Defoliation and below-ground herbivory in the grass Muhlenbergiaquadridentata : Effects on plant performance and on the root-feeder Phyllophaga sp. (Coleoptera, Melolonthidae)

In this study we evaluated (1) the combined effects of simulated defoliation and below-ground herbivory (BGH) on the biomass and nitrogen content of tillers and roots of the bunchgrass Muhlenbergia quadridentata and (2) the effect of defoliation on the survival of third-instar root-feeder larvae of Phyllophaga sp. The experiment was performed in a pine forest area at an altitude of 3200 m above sea level. The grass and the root-feeder species were native and dominant in the understory and in the macroarthropod root-feeder communities, respectively. Plants were established in pots in the field and were subjected to the following treatments in a factorial design: simulated defoliation (three levels) and BGH (with or without root-feeder larvae) with ten replicates per treatment. Plants were defoliated three times at 2-month intervals. The interaction between defoliation and root herbivory was significant for all components of plant biomass. In every case, light defoliation with BGH decreased live above-ground, root and total plant biomass, and the number of live tillers by more than 50% with respect to the same defoliation level without root-feeders. Plants apparently did not compensate for the carbon drain by root-feeders when a high proportion of older leaves were not removed by defoliation. Plants under heavy defoliation were not affected by the presence of root-feeders and showed a greater live/dead above-ground biomass ratio than lightly defoliated and control plants. Defoliation and BGH did not change tiller and root N concentrations but root herbivores did decrease live-tiller N content in lightly defoliated plants. Root-feeders but not defoliation decreased the root/shoot ratio by 40% and the live/dead above-ground biomass ratio by 45% through increased tiller mortality. Survivorship and final biomass of Phyllophaga sp. larvae were not affected by defoliation treatments during the 6-month study period. Received: 17 May 1996 / Accepted: 1 November 1996     

Attack dynamics of the pine shoot beetle, Tomicus piniperda (Col.; Scolytinae) in Scots pine stands defoliated by Bupalus piniaria (Lep.; Geometridae)

Abstract 1 After a 1‐year, extensive pine looper (Bupalus piniaria) outbreak, plots were laid out to study tree susceptibility to attack, and performance of Tomicus piniperda in pine trees suffering from varying levels of defoliation. 2 Tomicus piniperda was the dominating stem‐attacking species among the primary stem colonizers, and 82% of all trees that died had been colonized by T. piniperda. 3 Beetle attacks primarily struck severely defoliated trees, i.e. trees that suffered from 90% to 100% defoliation. 4 Beetle attacks peaked in the second year after cessation of the outbreak, and suppressed trees were both more frequently attacked and more susceptible to beetle attack than intermediate and dominant trees. 5 Trees surviving beetle attacks carried more foliage than trees that did not survive the attacks. 6 A single year of severe defoliation is enough to render pine trees susceptible to secondary pests, such as T. piniperda.