Host-plant acceptance on mineral soil and humus by the pine weevil Hylobius abietis (L.)

1 The pine weevil Hylobius abietis (L.) (Coleoptera, Curculionidae) is an economically important pest of conifer forest regeneration in Europe and Asia. 2 Soil scarification, which usually exposes mineral soil, is widely used to protect seedlings from weevil attack. However, the mechanism behind this protective effect is not yet fully understood. 3 Field experiments were conducted to determine the pine weevil's responses to visual and odour stimuli from seedlings when moving on mineral soil and on undisturbed humus surface. 4 One experiment measured the number of pine weevils approaching seedlings, with and without added host odour, on mineral soil and undisturbed humus. Seedlings with added host odour attracted more weevils on both soil types. Unexpectedly, somewhat more weevils approached seedlings surrounded by mineral soil. 5 In a similar experiment, feeding attacks on seedlings planted directly in the soil were recorded. Only half as many seedlings were attacked on mineral soil as on undisturbed humus. 6 In the first experiment, the weevils were trapped 2.5 cm from the bases of the seedlings' stems, whereas they could reach the seedlings in the experiment where seedlings were planted directly in the soil. We conclude that the pine weevils' decision on whether or not to feed on a seedling is strongly influenced by the surrounding soil type and that this decision is taken in the close vicinity of the seedling. The presence of pure mineral soil around the seedling strongly reduces the likelihood that an approaching pine weevil will feed on it.     

The effect of red wood ant abundance on feeding damage by the pine weevil Hylobius abietis

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Olfactory and visual stimuli used in orientation to conifer seedlings by the pine weevil, Hylobius abietis

Abstract. The influence of noncontact plant cues is investigated on the likelihood that individual conifer seedlings will be found by walking adults of the pine weevil, Hylobius abietis, in the field. Traps with solely odour or solely visual stimuli catch significantly more weevils than stimulus-free traps, and traps with the combination of odour and visual stimuli catch more weevils than traps with odour or visual stimuli alone. There is essentially an additive effect between odour and visual stimuli. The reactions to odour and visual stimuli are similar for three phases of the pine weevil's life cycle associated with three ages of clear-cuttings (i.e. sites where all trees have been harvested). Visual stimuli appear to be at least as important as odour for the pine weevil in finding an undamaged conifer seedling.     

Interaction between flight,reproductive development and oviposition in the pine weevil Hylobius abietis

• 1 The development of reproductive and flight capacity of pine weevils Hylobius abietis during the spring and their dispersal to, and subsequent development at, new clearfell oviposition sites comprise key phases in their life cycle in managed forests. At an old clearfell site where autumn‐emerging weevils had overwintered, weevils were trapped as they re‐emerged in the spring and tested for their ability to fly and then dissected to determine the degree of wing muscle and egg development.
• 2 Re‐emerging weevils were most abundant in pine growing at the edge of the clearfell and, over most of the trapping period (April to June), their capacity for flight (proportion flying and wing muscle width) was more advanced than in weevils from the clearfell itself, with a similar trend in the degree of reproductive development (proportion with mature eggs and egg volume).
• 3 In weevils from the clearfell, flight capacity and reproductive development increased concurrently to a peak around mid‐May. In weevils from pine, wing muscles were already well developed at the start of trapping, although few of them flew. Their more advanced development was attributed to the increased opportunities for maturation feeding after emergence in the previous autumn.
• 4 In the spring, weevils reached the canopy of trees for maturation feeding by walking and, to a lesser extent, by flight. Weevils dispersed by flight to oviposition sites in mid‐May when most of them were reproductively mature. After arrival, flight ability and wing muscle size declined rapidly but egg production was maintained until most weevils had stopped flying. When wing muscles reached their minimum size, there was a marked decline in egg size, suggesting that wing muscle breakdown is important in maintaining egg production at oviposition sites. Prospects for further wing muscle and reproductive development are discussed.

     

Movement behaviour of the pine weevil Hylobius abietis in relation to soil type: an arena experiment

Pine weevil, Hylobius abietis (L.) (Coleoptera: Curculionidae), movement behaviour on different substrates was studied in an indoor arena using a video camera and digital image processing technique. We analysed individual variation in movement characteristics, i.e. turning angles, movement directions, and distance moved per unit time on the bare and level arena surface which consisted of mineral soil (sand) and/or humus sections in various spatial configurations. Pine weevils on humus did not turn back when they came to the border with the sand. However, most individuals moved faster on sand than on humus. Thus, the results suggest that interactions between substrate differences and individual movement behaviour may to some extent explain why pine weevils have been observed to feed less frequently on coniferous seedlings planted on mineral soil than on those planted on humus.     

Alterations of the resin canal system of <Emphasis Type="Italic">Pinus pinaster</Emphasis> seedlings after fertilization of a healthy and of a <Emphasis Type="Italic">Hylobius abietis</Emphasis> attacked stand

Changes in resource availability and biotic and abiotic stress may alter the defensive mechanisms of pine trees. The effect of fertilisation on the resin canal structure of Pinus pinaster seedlings established in two trials in NW Spain, one attacked by Hylobius abietis and the other non-attacked, was studied. The leaders of 50 plants were destructively sampled and the resin canal density, the canal area and its relative conductive area in the phloem and xylem were assessed. Experimentally increased nutrient availability significantly decreased resin canal density in the phloem of the seedlings in the two analysed trials, where unfertilised seedlings presented up to 30% more resin canal density than the fertilised seedlings (mean value ± SEM = 0.32 ± 0.02 resin canals mm⁻² in the fertilised plants versus 0.45 ± 0.04 resin canals mm⁻² in the control plants). Fertilisation had no effect on the resin canal system in the xylem, but significantly increased tracheid size. Significant differences of resin canals among sites were observed mainly in the xylem; the resin canal density was 1.7-fold greater in the attacked site than in the non-attacked site. The similar structure of phloem resin canals in both sites supports that phloem resin canals are constitutive mechanisms of defence in P. pinaster, whereas xylem resin canals would be constitutive mechanisms but also inducible mechanisms of resistance following the attack of pine weevils or bark beetles. XM and LS equally contributed to this paper.     

Cues For Shelter Use in a Phytophagous Insect

Many insects spend a large proportion of their life inactive, often hiding in shelters. The presence of shelters may, therefore, influence where insects feed. This study examines stimuli affecting the use of shelters by adults of the pine weevil, Hylobius abietis (L.) (Coleoptera, Curculionidae). This species is an economically important forest pest in Europe since the adults feed on the stem bark of newly planted conifer seedlings. When there are hiding or burrowing places present in close proximity to a seedling, pine weevils may hide there and repeatedly return to feed on the same seedling. Experiments were conducted in a laboratory arena with above-ground or below-ground shelters and in the presence or absence of wind. Pine weevils were highly attracted to shelters both above and below ground. Weevils in shelters were often observed assuming a characteristic “resting” posture. Experiments with opaque and transparent shelters showed that visual stimuli are used for orientation towards shelters and also increase the probability of an individual remaining in a shelter. The presence of wind increased the weevils’ propensity to use shelters both above and below ground. The present study indicates that shelters have a major influence on the behavior of the pine weevil and possible implications of the results are discussed.     

Orientation of the pine weevil Hylobius abietis to underground sources of host volatiles

Adults of Hylobius abietis (L.) (Coleoptera: Curculionidae) were found to locate conifer roots suitable for oviposition by utilizing host volatiles diffusing through the soil. Underground sources of host volatiles were presented to weevils in a laboratory bioassay. A cold-trapping condensate of Scots pine, Pinus sylvestris L., and fractions of it were tested. Various fractions containing host terpenes attracted weevils in the bioassay, but the complete pine condensate caused the highest response. Ethanol was also found to be attractive. Weevils caged underground in the absence of host material did not attract weevils on the surface.

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Zusammenfassung Rüsselkäfer (Hylobius abietis) können auf der Erdoberfläche die Lage im Boden verborgenen Brutmaterials feststellen und Wurzeln senkrecht grabend auffinden. Mit einer neu entwickelten Methode für Laborversuche, die sich auf die spezifische Reaktion des Eingrabens zum Brutmaterial gründet, wurden das Orientierungsverhalten der Käfer und die zum Brutmaterial führenden Geruchsstoffe des Wirtes studiert.Beide Geschlechter von H. abietis reagierten gleichartig mit Eingraben auf die Geruchsstoffe des Wirtes. In Wahlversuchen zwischen gleichwertigen Anlockungspunkten kam häufig Aggregation der Käfer an einem Punkt vor. Die stärkere Ansammlung an einzelnen Stellen stand im Zusammenhang mit dem Vorhandensein einer Erdröhre zur Quelle der Geruchsstoffe. In den folgenden Versuchen wurde deshalb den Tieren jeweils nur eine Geruchsstoffquelle angeboten. Weder Weibchen noch Männchen im Boden übten eine Anziehung auf Käfer an der Oberfläche aus.Kiefernstücke und Kondensate flüchtiger Kieferinhaltsstoffe sowie Kondensat nach Passage durch die Kolonne des Gaschromatographen waren stark attraktiv. Alle durch präparative Gaschromatographie hergestellten Fraktionen des Kondensats waren ebenfalls, aber schwächer attraktiv. Zehnfache Verdünnung des Kondensats und der Fraktionen verminderte ihre Attraktivität mit etwa 40–70%. Auch Äthanol, Methanol und Pentan wurden geprüft; nur Äthanol hatte eine mässig anlockende Wirkung.

     

Diel behaviour and time budget of the adult pine weevil Hylobius abietis

The pine weevil [Hylobius abietis (L.); Coleoptera: Curculionidae] has a high economic impact on forest regeneration in Europe. The general biology of the pine weevil has received considerable attention, although there is insufficient knowledge about its diel behaviour and time budget. Therefore, in the present study, the feeding and locomotion behaviour of individual adult weevils on Norway spruce seedlings [Picea abies (L.) Karst.] is observed for 24‐h periods in the laboratory. Both girdled and nongirdled seedlings are used to assess how the behaviour of weevils is influenced by the physiological response of plants to the girdling. The locomotion pattern shows a distinct maximum during the beginning of the dark phase, whereas most feeding occurs during the second half of the dark phase and the first hours of the subsequent light phase. The girdling treatment increases the time that weevils spend on the seedlings during the first part of an observation session, although it has no effect on their feeding pattern. The time budgets of weevils on girdled and nongirdled seedlings are similar. On average, weevils spend 34% of their time in locomotion and 6% on feeding. Females spend more time feeding than males (7.1% versus 4.2%), possibly because they have higher food requirements (e.g. for egg production). Females also spend more time in total on the seedlings than males (26.3% versus 7.0%). The present study reveals, in high temporal resolution, the diel feeding and locomotion behaviour and time budget of male and female pine weevils.     

Growth and nutritional response of Pinus pinaster after a large pine weevil (Hylobius abietis) attack

Hylobius abietis is an important pest of coniferous plantations in Europe, to which high mortality, stem deformities, and growth loss are typically attributed. In pine trees, as in other long-lived organisms, there is uncertainty regarding the long-term costs of short-term resistance against invading organisms. We examined the nutritional status of Pinus pinaster after a 2-year long H. abietis attack, measuring needle and phloem N and P concentrations, and the impact of the damage on subsequent growth, survival, and stem deformities over a period of 5 years. The study sites were a P. pinaster family × fertilization trial, and a neighbouring twin trial with similar climate and soil characteristics that was not attacked. Growth losses after the H. abietis attack were important (up to 40%), but restricted to the first years after the attack. Five years after the attack, the annual height increment of pines in the attacked stand was not related to the initial damage suffered, and plants showed regular stems, normal leader dominance, and regular height after 5 years. These findings suggest strong compensatory growth in P. pinaster and indicate relatively high tolerance to the large pine weevil. Needle nutrient concentrations in the healthy stand were, as expected, significantly greater in experimentally fertilized plants, and they were linearly related to those in phloem showing equilibrated stoichiometry both for nitrogen (r = 0.86; P < 0.01; N = 25) and phosphorus (r = 0.84; P < 0.01; N = 25). However, at the attacked stand, nutrient concentrations in the needles did not follow the experimentally manipulated nutrient availability in soils, and phosphorus concentration in the needles was unexpectedly not related to those in the phloem. The pine seedlings attacked by H. abietis showed altered potential of allocating nutrients to their tissues according to the nutrient availability existing in the soil, as well as altered stoichiometry in N and P concentrations among phloem and leaves. Maritime pine seems to be tolerant to the pine weevil attack, at least in the conditions of this study, where pine weevil damage caused a deep alteration of nutrient allocation and nutritional status. Further research is needed to elucidate to what extent altered nutrient allocation may be part of an induced response to the attack or just derived from the vascular injury caused by the weevil wounding in the phloem.     

The large pine weevil Hylobius abietis (L.) as a potential vector of the pathogenic fungus Diplodia sapinea (Fr.) Fuckel

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Effects of green tree retention, prescribed burning and soil treatment on pine weevil (Hylobius abietis and Hylobius pinastri) damage to planted Scots pine seedlings

1 Feeding damage and mortality caused to planted Scots pine seedlings by the pine weevils Hylobius abietis and Hylobius pinastri were studied on burned and unburned sites with 0, 10 and 50 m³ per hectare levels of green tree retention from the second to the fourth summer after logging and burning of the sites. 2 The rate of severe feeding damage to pine seedlings caused by pine weevils was higher on burned clearcut sites than on unburned ones, whereas burning did not increase the feeding damage rate on sites with groups of retention trees. The damage rate in the fourth summer was approximately the same on burned and unburned sites. 3 Pine weevil feeding was the major cause of mortality of freshly planted pine seedlings on unburned sites. On burned sites, mortality was higher than the rate of severe feeding damage, particularly in the second summer after burning, possibly owing to fungal attack and abiotic factors. 4 At a retention tree level of 50 m³, feeding damage to the seedlings was lower than on clearcuts and at a 10 m³ retention tree level. Furthermore, on sites with 50 m³ of retention trees, scarification of the soil was found to decrease feeding damage more effectively than on clearcuts and 10 m³ sites. If the seedlings were situated in the centre of scarified patches, scarification alone was as effective as insecticide treatment on unscarified soil for decreasing feeding damage and mortality. 5 The results suggest that when burning is applied as silvicultural treatment after clear‐cuts, retention of trees is recommended to reduce the damages caused by pine weevils on pine seedlings.     

Pine weevil damage to Norway spruce seedlings: effects of nutrient‐loading,soil inversion and physical protection during seedling establishment

• 1 The pine weevil Hylobius abietis (L.) feeds on the bark of young conifer seedlings and is one of the most economically important forest pests in Europe.
• 2 In a field experiment, we examined the combined effects of the treatments: nutrient‐loading of seedlings, planting in scarified plots and protection of seedlings against pine weevil damage for either half a season or a full season.
• 3 Nutrient loading had no significant effect on the amount of pine weevil feeding.
• 4 Seedling mortality was significantly reduced when seedlings were protected from pine weevil feeding during establishment. This occurred even though the debarked area of protected seedlings 5 weeks after the shields had been removed was similar to that of the unprotected seedlings. This indicates that initial protection rendered the seedlings more tolerant to later wounding by pine weevil.
• 5 Planting in soil inversion significantly reduced feeding compared with planting in humus.
• 6 We conclude that nutrient‐loading of seedlings in the autumn before planting would not increase pine weevil feeding after planting. Mortality could be reduced by treatments that postpone the start of pine weevil feeding on seedlings. Such treatments, combined with planting in soil inversion, would result in increased seedling growth, vitality and tolerance to pine weevil attack.

     

Effects of green tree retention, prescribed burning and soil treatment on pine weevil (Hylobius abietis and Hylobius pinastri) damage to planted Scots pine seedlings

1 Feeding damage and mortality caused to planted Scots pine seedlings by the pine weevils Hylobius abietis and Hylobius pinastri were studied on burned and unburned sites with 0, 10 and 50 m³ per hectare levels of green tree retention from the second to the fourth summer after logging and burning of the sites. 2 The rate of severe feeding damage to pine seedlings caused by pine weevils was higher on burned clearcut sites than on unburned ones, whereas burning did not increase the feeding damage rate on sites with groups of retention trees. The damage rate in the fourth summer was approximately the same on burned and unburned sites. 3 Pine weevil feeding was the major cause of mortality of freshly planted pine seedlings on unburned sites. On burned sites, mortality was higher than the rate of severe feeding damage, particularly in the second summer after burning, possibly owing to fungal attack and abiotic factors. 4 At a retention tree level of 50 m³, feeding damage to the seedlings was lower than on clearcuts and at a 10 m³ retention tree level. Furthermore, on sites with 50 m³ of retention trees, scarification of the soil was found to decrease feeding damage more effectively than on clearcuts and 10 m³ sites. If the seedlings were situated in the centre of scarified patches, scarification alone was as effective as insecticide treatment on unscarified soil for decreasing feeding damage and mortality. 5 The results suggest that when burning is applied as silvicultural treatment after clear‐cuts, retention of trees is recommended to reduce the damages caused by pine weevils on pine seedlings.     

Ants protect conifer seedlings from feeding damage by the pine weevil Hylobius abietis

• 1 Ants that protect food resources on plants may prey on (or deter) herbivores and thereby reduce damage. Red wood ants (of the Formica rufa group) are dominant ants in boreal forests of Eurasia and affect the local abundance of several herbivorous species.
• 2 The pine weevil Hylobius abietis (L.) is a herbivore that causes severe damage by feeding on the bark of coniferous seedlings within areas of forest regeneration.
• 3 We investigated whether ants can protect conifer seedlings from pine weevil feeding. In a manipulative experiment, ants were attracted to sugar baits attached to spruce seedlings and the damage caused by pine weevils was compared with control seedlings without ant‐baits.
• 4 The feeding‐scar area was approximately one‐third lower on the seedlings with ant‐baits compared with the controls. Besides red wood ants, Myrmica ants were also attracted in high numbers to the ant baits and the relative effects of these species are discussed.
• 5 The results obtained in the present study support the trophic cascade hypothesis (i.e. damage to herbivores is suppressed in the presence of predators). The decreased pine weevil feeding on the baited seedlings was probably a result of nonconsumptive interactions [i.e. the presence of (or harassment by) ants distracting pine weevils from feeding].
• 6 Understanding the role of ants may have important implications for future strategies aiming to control pine weevil damage. For example, maintaining suitable conditions for ants after harvesting stands may be an environmentally friendly but currently unexploited method of for decreasing weevil damage.

     

Lignosulfonate promotes the interaction between Scots pine and an ectomycorrhizal fungus Pisolithus tinctorius in vitro

Lignosulfonate (LS) is a lignin-based polymer obtained as a by-product from paper industry, which may have potential as an amendment with macronutrients. We studied effects of LS on the interaction between Scots pine (Pinus sylvestris L.) seedlings and hypocotyl cuttings and the ectomycorrhizal (ECM) fungusPisolithus tinctorius (Pers.) Coker and Couch. The experiments were performed in vitroon the MMN agar medium containing Fe–LS chelate at the concentrations of 0, 5, 10 and 25 mg/L. Inoculation with P. tinctoriusincreased root growth of the seedlings. Fe–LS enhanced P. tinctorius induced formation of lateral roots and had a dose-dependent positive effect on the establishment of mycorrhizas on the seedlings. The growth of the fungal mycelium was improved by Fe–LS, which might cause faster and more intensive contact with the roots and, thus, better root growth and mycorrhiza formation. P.tinctorius enhanced also adventitious root formation and subsequent root growth of the hypocotyl cuttings but without any synergistic effect with Fe–LS. Our study with P. tinctorius and Scots pine in vitro indicates that a low-cost by-product Fe–LS, obtained from paper industry, may be a potential tool to improve the efficiency of fungal inoculations, thus, facilitating the early interaction between an ECM fungus and host seedling.     

Abscisic acid in shoots and roots of Scots pine (Pinus sylvestris L.) seedlings grown in controlled long-day and short-day environments

Scots pine (Pinus sylvestris L.) seedlings grown in nutrient solution in controlled-environment chambers were used. The effects of a shortday (SD, early autumn) treatment on growth and the content of free and alkaline hydrolysable abscisic acid (ABA) in shoots and roots were investigated. The weekly relative growth rates of seedlings grown continuously under long-day (LD, summer) conditions were stable at approx. 0.08 g g^–1 d^–1 between weeks four and eight from germination. Weekly relative growth rates of seedlings transferred to SD conditions decreased rapidly to a then stable level of approx. 0.04 g g^–1 d⁰¹. Shoot elongation ceased within two weeks of SD treatment. The content of both free and alkaline hydrolysable ABA was approx. 40–50% higher in shoots of seedlings grown for five weeks in LD plus one week in SD than in shoots of seedlings grown for five or six weeks in LD. Two additional weeks of SD did not change the free ABA content. Three weeks in simulated late autumn (SD but decreased temperatures) and three weeks in simulated winter (lower light intensity and temperature) further increased the content of free ABA in the shoots. A transfer back to LD conditions reduced the ABA content to a level equal to the level found during the first LD period. The recovery of radioactive ABA at certain times after application ofr[³H] ABA was the same in shoots and roots of LD-grown and SD-treated seedlings.Abbreviations ABA abscisic acid - LD long day(s) - RGR₇ weekly relative growth rates - SD short day(s)     

Fatty acids,starch and biomass of Scots pine needles and roots in open-air ozone exposure

Scots pine (Pinus sylvestris L.) seedlings were fumigated with 1.2–1.5 x ambient ozone (cumulative exposure) over 2 seasons in an open-air experiment. Starch and fatty acid concentrations were analyzed in needle and root tissue in the summer, autumn and early winter. Seedling growth was determined by measuring the height of the stem and the total shoot and root biomass. Significant decreases in growth were found in exposed seedlings, even though visible symptoms were lacking. Almost significant reductions in needle and root starch concentrations were found. In the ozone treated foliage, significant increases in myristic acid (140) were detected, but the major fatty acids remained unchanged. Fatty acid ratios showed that the degree of unsaturation decreased in treated needles in the summer. In the roots of ozone treated seedlings, changes in fatty acids were different from those in the foliage. Decreases of the main root fatty acids (160, 180, 181, 18:2, 183) were detected in the summer. These results show that Scots pine is susceptible to enhanced levels of ozone. If the tropospheric ozone levels continue to increase it may have deleterious effects on Scots pine forests in Finland.     

Non-pesticide alternatives for reducing feeding damage caused by the large pine weevil (Hylobius abietis L.)

The large pine weevil (Hylobius abietis L.) is an important pest of young forest stands in Europe. Larvae develop under the bark of freshly cut pine and spruce stumps, but maturing weevils feed on the bark of coniferous seedlings. Such seedlings frequently die because of bark consumption near the root collar. We tested the effect of three treatments (the insecticide alpha cypermethrin, a wax coating and a glue coating) on the feeding damage caused by H. abietis on Douglas fir (Pseudotsuga menziesii) and Norway spruce (Picea abies) seedlings under semi-natural conditions. In two experiments (one in 2016 and another in 2017) seedlings in cages were subjected to pine weevil feeding for 16 weeks under shaded outdoor conditions. The experiment in 2016 compared insecticide and wax treatments and an untreated control on Douglas fir and Norway spruce, and the experiment in 2017 compared insecticide, wax and glue treatments and an untreated control on Norway spruce. In both experiments, all treatments significantly reduced H. abietis feeding damage at week 8 at the end of both experiments (week 16); the effect of treatments was significant only on spruce seedlings. The damages on Douglas fir seedlings was less on treated seedlings than on untreated control seedlings but differences were not significant. Coating stems with glue and especially with wax was generally effective at reducing weevil damage and in most cases provided control that was not significantly different from that provided by insecticide treatment. Our results suggest that a wax coating has the potential to replace the protection of seedlings provided by insecticides.     

Factors affecting seedling recruitment and survivorship of the Japanese subalpine stone pine, <Emphasis Type="Italic">Pinus pumila</Emphasis>, after seed dispersal by nutcrackers

To clarify the factors affecting the seedling establishment of the subalpine stone pine (Pinus pumila Regl.) from nutcracker-cached seeds, subsequently emerged seedlings were examined in two plots on Mt Akitakomagadake, northern Japan. The survivorship of older seedlings (2–20years old) was also monitored during six seasons at nine different sites. In one plot with 18 caches, seed germination occurred between June and mid-July. During this period, nutcrackers retrieved the stored pine seeds from 16 caches and ate some seeds immediately. Simultaneously, this nutcracker behavior caused mechanical damage to newly emerging seedlings (e.g. uprooting and tearing off cotyledons). Such initial loss to in situ harvesting and mechanical damage accounted for 75% of the total seeds remaining in the caches. The number of established current seedlings sharply declined during the first 2years, and the survival rate was 4.4% over four winters. Two major mortality factors were identified: uprooting by frost-heave soil disturbance in the spring of the second year and standing death by drought or other physiological stresses in early summer. In another plot with 13 caches, survivorship of newly emerged seedlings was also low (4.2%), but mortality was mostly due to summer drought, indicating that the frost-heave event was a site-specific disturbance factor. For older seedlings, survival rates reached approximately 90% even after six seasons and summer drought stress was a major mortality factor. My findings suggest that seedling recruitment of P.pumila was largely limited by both nutcracker disturbance and external disturbance (and/or stresses) at an early stage. However, after the critical first few years, pine seedlings were highly likely to survive and grow to the sapling stage despite the harsh environment of the high mountains.