Effects of host plant hybridization on resistance to willow leaf rust caused by Melampsora sp.

Patterns of resistance to willow leaf rust caused by Melampsora sp. were measured on Salix eriocephala, Salix sericea, and their interspecific hybrids in natural field populations in 1991–94 and on potted plants in common garden experiments in 1994 and 1995. Testing was carried out to establish if the resistance of hybrids was intermediate between pure parental species (additive), equal to one of the parents (dominant), lower than both parents (susceptible), higher than both parents (resistant), or not different than either parent (null hypothesis). Support was generally found for hybrid susceptibility in the field and common garden experiments, with the null hypothesis of no difference having some support. The results from the common garden study suggest that the susceptibility of hybrids seen in the field is due to genetic differences. Hybridization between these species of willows results in increased susceptibility to this pathogen. The variation among years in the relative susceptibility of hybrid and parental willows may be due to environmental effects on the willows or to shifts in the abundance of Melampsora races.     

Long term losses caused by foliar diseases on growth and survival of Eucalyptus globulus in Uruguay

Eucalyptus globulus is the most important forest species in Uruguay, with more than 250,000 ha of commercial plantations. Despite its high susceptibility to diseases, production losses caused by foliar diseases have not been properly quantified in this country. This study analyzes the effects of foliar damage on growth and survival using data from a progeny test of E. globulus naturally infected by Teratosphaeria leaf disease and eucalypt rust (Puccinia psidii). The severity of leaf spots and defoliation were quantified 8 months after planting and tree growth and mortality were evaluated 2, 4 and 6 years later. The trial had a high incidence of foliar damage, with a mean leaf spot severity of 28.7% and a mean defoliation of 37%. The greatest impact of foliar damage, both on growth rate and mortality, occurred in the first 2 years after damage was assessed. During this period, leaf spot severity less than 40% and defoliation below 50% did not affect growth, while survival was affected when leaf damage was 70% or greater. By the sixth year both stem growth and survival were affected by severe foliar damage (spotting or defoliation of 80% or more), with a loss of up to 25% in diameter and an accumulated mortality over 70%. It has been established for the first time that under the intensive Uruguayan productive conditions, E. globulus trees can tolerate a relatively high degree of leaf spotting or defoliation but severe foliar damage in the first months can cause considerable production losses, putting at risk the economical viability of this species.     

Condensed tannins in Leucaena and their relation to psyllid resistance

Leucaena species and hybrids were examined to determine the relative content of condensed tannins and its relationship to susceptibility to defoliation by the Leucaena psyllid (Heteropsylla cubana). The interspecific Leucaena hybrid KX2 (L. leucocephala × L. pallida) was examined for variability of condensed tannin levels. A broad range of condensed tannin values were found which indicate potential for improved forage value through selection for KX2 trees with lower condensed tannin while maintaining psyllid resistance. A significant correlation was not found linking condensed tannins and psyllid resistance.     

Growth, cold-hardiness, protein content, and digestibility of 70 Leucaena seedlots on three sites in Texas, USA

Seventy seedlots of Leucaena leucocephala, L. pallida, L. diversifolia, L. retusa, L. esculenta, L. confertifolia, L. greggii and L. pulverulenta and various hybrid combinations were examined for survival to freezing weather, biomass production, leaf protein content and in vitro dry matter digestibility in Texas. Three sites along a 600 km north-south transect were chosen to provide a wide range in exposure to freezing weather. The minimum temperature on the warmer Three Rivers site was −7 °C while the minimum temperatures on the colder Vernon site was −14 °C. At the Three Rivers site where −7 °C temperatures occurred and where forage production was a priority and some winter frost damage was not a problem, L. leucocephala K636 and hybrids containing L. diversifolia and L. pallida have the most promise. If total lack of damage to −7 °C is essential, species such as L. retusa, L. pulverulenta (seedlot 61, 62) and hybrids such as L. retusa × L. greggii (53) and L. retusa × L. pulverulenta (55) have the greatest potential. This study examined only one family from each of the cold-hardy species as a parent in the promising hybrids. As extensive intraspecific genetic variation is present within these leucaena species there is great potential to find hybrids with much better combinations of cold hardiness, forage production, leaf protein and dry matter digestibility. This revised version was published online in June 2006 with corrections to the Cover Date.     

Responses of Prunus ferganensis,Prunus persica and two interspecific hybrids to moderate drought stress

Prunus ferganensis (Kost. & Riab) Kov. & Kost, a close relative of cultivated peach (Prunus persica (L.) Batsch.), is native to arid regions of central Asia. A distinguishing feature of P. ferganensis is its prominent, elongated, unbranched pattern of leaf venation. To determine whether the long-vein trait could be used as a marker in breeding for drought tolerance, we investigated the association between this trait and the leaf morphological and physiological parameters related to drought response in P. ferganensis, P. persica and two interspecific hybrids, one with the long-vein trait (BY94P7585) and one without (BY94P7589). The four genotypes were grafted onto "Guardian" peach rootstock and half of the plants were assigned to a drought treatment in which irrigation was limited to 25-50% of the evapotranspiration (ET) rate measured in the remaining well-watered plants, which were irrigated to runoff daily. The drought treatment reduced photosynthesis and leaf conductance by 49-83% and reduced total leaf area per plant by 17-24%, but generally did not affect mid-morning leaf water potential. Leaf gas exchange did not differ among genotypes in either treatment. Sorbitol accumulated in mature leaves in response to drought, but neither its amount nor its metabolism varied systematically with climatic adaptation among genotypes. Accumulation of transport sugars was highest in P. ferganensis, indicating that growth reduction may represent an important strategy for coping with drought in this species. Prunus ferganensis and the hybrids had higher ET than P. persica, and seemed to use water opportunistically, maintaining high gas exchange rates and consequently high ET when water was available, and avoiding low water potentials through stomatal closure as soil water declined. Leaf size (cm(2) leaf(-1)) and specific leaf area (cm(2) g(-1) dry mass) were lower in P. ferganensis and the hybrids than in P. persica. We conclude that the long-vein trait is not a reliable marker for drought tolerance, but leaf traits of P. ferganensis such as size and specific leaf area may be useful in P. persica breeding programs targeting drought tolerance.     

Ophiostoma ulmi metabolites and elm cell membrane permeability. Possible use in early tests of resistance

Electrolyte leakage from leaf discs of three Ulmus species when challenged with cultural filtrate of an NAN aggressive isolate of Ophiostoma ulmi paralleled their susceptibility to the disease in the field. But the correlation was not working with some hybrids derived from the common field elm showing intermediate resistance. The possible explanation will be checked in further tests which are in progress to assess the possibility of using this technique in early screening for resistance.     

Susceptibility of winter‐dormant Pinus sylvestris families to vole damage

Two‐ and eight‐year‐old Pinus sylvestris, originating as controlled crossings from southern and northern Finland, were studied in order to determine their susceptibility to damage by Microtus‐species. For characteristics that indicate climatic adaptedness (bud set, needle stage) there was a clinal change from north to south according to origin. These differences were not correlated with susceptibility to vole damage. Nor was the total phenol content of the saplings correlated with resistance to such damage. Between families, however, there were statistically significant differences in susceptibility to damage. Some families showed both rapid rate of growth and high resistance.     

Susceptibility of Corymbia species and hybrids to arthropod herbivory inAustralian subtropical hardwood plantations

Corymbia hybrids are becoming significant plantation varieties in subtropical and tropical Australian plantation forestry. Although primarily developed for disease resistance and amenability to clonal propagation, they have also proven to have good growth rates and site plasticity. Here we examined the susceptibility of pure Corymbia species and hybrids to pest attack. Three trial sites containing C. citriodora subsp. variegata, C. torelliana, and the hybrids C. torelliana × C. citriodora subsp. variegata, C. torelliana × C. citriodora subsp. citriodora and C. torelliana × C. henryi were assessed for pest identity, incidence and severity. Pests caused about three-quarters of the visible crown damage to trees in these trials. At the site that had the most arthropod damage, hybrid trees had higher damage scores and higher growth scores (height, diameter at breast height over bark, and volume) than pure species. Site was more important than taxon in explaining damage scores, and taxa performed differently for most traits between sites. Tree growth was negatively correlated with general crown damage, while arthropod damage alone showed no significant relationship with growth. Our results highlight the importance of establishing taxa trials across a range of sites when selecting for pest resistance.     

Variation components in leaf morphology of recruits of two hybridising oaks [<Emphasis Type="Italic">Q. petraea</Emphasis> (Matt.) Liebl. and <Emphasis Type="Italic">Q. pyrenaica</Emphasis> Willd.] at small spatial scale

Leaf morphological variation was examined in recruits of two hybridising oaks in a small sympatric area from Central Spain. Nuclear microsatellites were used to identify hybrids and assess the parental lineage. By Bayesian clustering analysis, 5% of hybrids were found. Principal component analysis was used to reduce 15 morphometric variables to four components associated with leaf size, lobation/pubescence and overall shape of the leaf. The percentage of variance due to genetic factors was evaluated through nested analysis of variance. As much as 70% of variance component was due to the factor “species” for lobation/pubescence, suggesting high adaptive value for these traits, possibly related to ecological constraints of the species. The genetic component of variance for leaf size and overall shape of the leaf was below 33%. Age and height of the recruits did not correlate with sun-leaf morphology. Competition indexes and diameter of the recruits showed slight, although significant, correlations with leaf size and lobation/pubescence components, pointing to some trade-offs between competition for light and leaf morphology of Q. petraea and Q. pyrenaica recruits.     

Water relations and gas exchange of Acer saccharum seedlings in contrasting natural light and water regimes

Field measurements were made of leaf photosynthesis (A), stomatal conductance (g) and leaf water relations for sugar maple (Acer saccharum Marsh.) seedlings growing in a forest understory, small gap or large clearing habitat in southwestern Wisconsin, USA. Predawn water status, leaf gas exchange and plasticity in field and laboratory water relations characteristics were compared among contrasting light environments in a wet year (1987) and a dry year (1988) to evaluate possible interactions between light and water availability in these habitats. Leaf water potentials (Psi(leaf)) at predawn and midday were lower for clearing than gap or understory seedlings. Acclimation of tissue osmotic potentials to light environment was observed among habitats but did not occur within any of the habitats in response to prolonged drought. During a summer drought in 1988, decreases in daily maximum g (g(max)) and maximum A (A(max)) in clearing seedlings were correlated with predawn Psi(leaf), which reached a seasonal minimum of -2.0 MPa. Under well-watered conditions, diurnal fluctuations in Psi(leaf) of up to 2.0 MPa in clearing seedlings occurred along with large midday depressions of A and g. In a wet year, strong stomatal responses to leaf-to-air vapor pressure difference (VPD) in sunny habitats were observed over nine diurnal courses of gas exchange measurements on seedlings in a gap and a clearing. Increasing stomatal limitations to photosynthesis appeared to be responsible for the reduction in A at high VPD for clearing seedlings. In understory seedlings, however, low water-use efficiency and development of leaf water deficits in sunflecks was related to reduced stomatal limitations to photosynthesis relative to seedlings in sunny habitats. Predawn Psi(leaf) and VPD appear to be important factors limiting carbon assimilation in sugar maple seedlings in light-saturating irradiances, primarily through stomatal closure. The overall results are consistent with the idea that sugar maple seedlings exhibit "conservative" water use patterns and have low drought tolerance. Leaf water relations and patterns of water use should be considered in studies of acclimation and species photosynthetic performance in contrasting light environments.     

Soil properties and tree growth on rehabilitated forest landings in the interior cedar hemlock biogeoclimatic zone: British Columbia

We studied operational landing rehabilitation programs in three forest districts of interior British Columbia (BC). Winged subsoiling and grass/legume seeding, followed by planting of lodgepole pine (Pinus contorta var. Latifolia) generally resulted in successful re-establishment of forest cover on landings. In the Boundary district, fifth year tree heights on landings were not significantly different from those in adjacent plantations, and fifth year growth increments were also similar despite evidence of delayed seedling establishment caused by cattle grazing damage. Trees on landings in the Kalum district were shorter than those in plantations, but fifth year increments were similar. Landings in the Kalum with >20% clay had lower stocking densities, tree heights, and probe depths compared to landings with <20% clay content. In the Kispiox district, average fifth year tree heights and fifth year increments were lower on landings than those on plantations. In the Kispiox district, landings with >20% clay content had shorter trees growing on them when compared to tree heights on landings with <20% clay. Landings in the Kispiox had the least probe depth of the three districts, and the greatest difference in height and increments between landings and plantations, supporting field reports of poor decompaction effectiveness there. In all districts, there was no forest floor present on landings 7 years after subsoiling, and cover of non-coniferous vegetation was lower than for plantations. Higher soil temperatures and lower soil moisture contents were recorded on landings relative to plantations in all districts during the growing season of 1998. Landing soils also had lower total C, N and mineralizable-N (min-N) than plantation soils in the Boundary and Kispiox, but such differences were not statistically significant in the Kalum. Reduced levels of total C, N and min-N on landings did not appear to have affected current foliar nutrient status of trees growing on these sites, as most nutrients appeared in concentrations considered adequate or only slightly deficient, except for foliar S on all sites. Average concentrations of N, P, K and S in foliage were either similar to or significantly higher for trees on landings relative to plantations, but these differences were attributed to dilution in plantation foliage rather than deficiency. Our results support the conclusion that operationally feasible techniques for soil rehabilitation can create conditions suitable for establishment of a new forest on sites that otherwise would be considered non-productive. Stocking levels and tree growth rates observed in our study are consistent with the conclusion that a commercial forest may be produced on some of the rehabilitated areas.     

Testing of selected South African <Emphasis Type="Italic">Pinus</Emphasis> hybrids and families for tolerance to the pitch canker pathogen, <Emphasis Type="Italic">Fusarium circinatum</Emphasis>

Plantations of Pinus spp. constitute approximately 50% of the South African forestry industry. The first aim of this study was to develop a reliable inoculation technique to screen Pinus spp., for tolerance to infection by F. circinatum, which threatens pine forestry in South Africa. Inoculation of branches was compared with stem inoculations and we considered the number of branches or trees required to obtain statistically significant results. Furthermore, variation in the susceptibility of some Pinus families, clones and hybrids was considered. Results showed that branch inoculations were closely correlated with those from stem inoculations, and that it is important to consider branch and stem diameters when assessing susceptibility of trees. Subsequent trials using branch inoculations showed significant differences in F. circinatum tolerance amongst a range of pine species and hybrids of potential interest to forestry in South Africa. Significant differences in susceptibility were also found among clones of two P. radiata families. The most tolerant trees were P. elliottii × caribaea and P. patula × oocarpa hybrids, while the most susceptible species were P. patula, P. greggii and hybrids of these two. This is the first trial considering the susceptibility of Pinus hybrids, Pinus clones and some P. patula provenances, and the results indicate excellent potential for breeding for tolerance to pitch canker in South Africa. Application The accurate selection of disease tolerant planting stock for the South African forestry industry is crucially important for the continued sustainability of this important industry. The work described here provides valuable information on an artificial inoculation technique that will assist the industry in screening trees for tolerance to the pitch canker fungus, F. circinatum. It also provides some indication of the relative susceptibility of a number of Pinus spp., hybrids and families currently being evaluated in the country.     

Ontogenetic changes in stomatal and biochemical limitations to photosynthesis of two co-occurring Mediterranean oaks differing in leaf life span

A quantitative analysis was applied to the stomatal and biochemical limitations to light-saturated net photosynthesis under optimal field conditions in mature trees and seedlings of the co-occurring evergreen oak, Quercus ilex L., and the deciduous oak, Q. faginea Lam. Stomatal limitation to photosynthesis, maximal Rubisco activity and electron transport rate were determined from assimilation versus intercellular leaf carbon dioxide concentration response curves of leaves that were subsequently analyzed for nitrogen (N) concentration, mass per unit area, thickness and percent internal air space. In both species, seedlings had a lower leaf mass per unit area, thickness and leaf N concentration than mature trees. The root system of seedlings during their third year after planting was dominated by a taproot. A lower leaf N concentration of seedlings was associated with lower maximal Rubisco activity and electron transport rate and with assimilation rates similar to or lower than those of mature trees, despite the higher stomatal conductances and potential photosynthetic nitrogen-use efficiencies of seedlings. Consequently, stomatal limitation to photosynthesis increased with tree age in both species. In both seedlings and mature trees, a lower assimilation rate in Q. ilex than in Q. faginea was associated with lower stomatal conductance, N allocation to photosynthetic functions, maximal Rubisco activity and electron transport rate, and potential photosynthetic nitrogen-use efficiency but greater leaf thickness and leaf mass per unit area. Tree-age-related changes differed quantitatively between species, and the characteristics of the two species were more similar in seedlings than in mature trees. Despite higher stomatal conductances, seedlings are more N limited than adult trees, which contributes to lower biochemical efficiency.     

Vegetative Propagation of Leucaena Hybrids by Cuttings

Many interspecific hybrids have been produced among Leucaena species, some with great potential for high-value wood and forage production. Superior hybrids are often highly sterile, and vegetative propagation would be essential for their exploitation. Four experiments were conducted to investigate factors affecting the rooting of cuttings of Leucaena hybrids. Nine hybrids were studied that represented different ploidy levels (2x, 3x, and 4x), different levels of seedlessness, different ages (5–16 years) and varying leaflet sizes. Rooting ability basically depended on genotype, with only a few hybrids failing to root. Cuttings required leaflets to root, and half or more of a leaf led to higher frequencies of better roots. One-node cuttings had the same rooting ability as two-node cuttings. Rooting ability decreased dramatically in the cooler, low-light winter seasons of lowland Hawaii. Etiolation treatments increased rooting ability significantly for two difficult-to-root hybrids.     

Leaf longevity of western red cedar (Thuja plicata) increases with depth in the canopy

Leaf longevity determines the annual cost of replacing foliage biomass and influences water and nutrient budgets. Longevity is readily estimated in most evergreen species by annual bud scars along the shoot. However, some species with indeterminate growth do not show these annual markers, making estimation of longevity difficult. One of these species is the widespread and economically valuable western red cedar (Thuja plicata J. Donn ex D. Don), for which no dependable estimates of leaf longevity exist. In this study, we estimated leaf longevity for western red cedar by counting growth rings in shoots at the point of leaf abscission. Estimates were obtained on 26 dominant or codominant trees growing in natural stands in a montane forest in northern Idaho, USA. Leaf longevity averaged 8.9 (SE = 0.2) years, but it strongly increased with depth in the canopy (0.3 year m(-1); mean crown depth was 15 m), increasing from a mean of 6.8 years in the upper third of the canopy to 10.6 years in the lower third. The increase in longevity with depth in the crown is consistent with many reports showing that longevity increases in resource-limiting environments. Longevity did not vary significantly with altitude or solar insolation in these montane forests. Among stand-level variables, longevity was correlated only with leaf area index: it increased slightly in stands with high leaf area indices. This approach to longevity estimation may be useful for any species that produces annual rings but no obvious bud scars, including many Cupressaceae species.     

Premature losses of leaf area in response to drought and insect herbivory through a leaf lifespan gradient

Implications of the differences in leaf life span are still subject to debate in the field of ecophysiology.Since leaf traits associated with these differences may be decisive for determining the distribution of tree species,this topic is particularly relevant in the context of climate change.This study analyzes the effects of the differences in leaf life span on premature losses of leaf area owing to insect herbivory and to abiotic stress.Loss of leaf area may be an important determinant of total leaf carbon assimilation.Seven Mediter-ranean tree species,distributed on four sites with different climates were studied.The species exhibited strong differ-ences in leaf life span and in leaf traits,especially leaf mass per unit area.Premature leaf area losses were estimated in response to insect herbivory and summer drought over two years.The results revealed that,despite having older leaf cohorts with more damage,species with longer leaf dura-tion had lower area lost to herbivores and less damage due to accelerated senescence during the summer drought.With respect to the predicted increase in water stress,deciduous species are at a disadvantage due to their high premature loss of leaf area and thus loss of photosynthetic capacity.     

Influence of soil water on the physiological and morphological components of plant water balance in Populus trichocarpa, Populus deltoides and their F(1) hybrids

Patterns of leaf growth, transpiration and whole-plant water balance in Populus trichocarpa, P. deltoides and their F(1) hybrids were studied during a soil drying cycle. Plant responses were analyzed during three distinct stages of dehydration. In stage I, the transpiration rate of drought-stressed plants remained constant and equal to that of well-watered plants even though soil water content declined by more than 40%. Stage II began as soil and plant water deficits induced stomatal closure. When soil water was expressed as a fraction of transpirable soil water, the transition from stage I to stage II occurred at soil water fractions of 0.35, 0.45 and 0.60 for P. trichocarpa, P. deltoides and their F(1) hybrids, respectively. Reductions in leaf growth coincided with the shift from stage I to stage II. As soil water declined further, decreases in relative transpiration and whole-plant leaf area were significantly greater in parental species than in F(1) hybrids. Inherent feedbacks controlling stomatal water loss and the maintenance and growth of leaf tissue appeared to differ between F(1) and parental genotypes in a pattern characteristic of an overdominant mode of inheritance.Stage III began once the ability of stomata to compensate for water loss had been exhausted. Substantial differences were found in plant survival during stage III, with F(1) hybrids surviving longer than parental species. Survival was more strongly correlated with the hydraulic conductivity of xylem tissues than with the dehydration tolerance of leaf tissues. Collectively, these responses suggest that F(1) hybrids were more drought resistant than either parental species and highlight the importance of whole-plant studies of functional relationships between plant growth, water balance and hydraulic conductivity.     

Wind Damage to the Leaves of Sycamore (Acer pseudoplatanus L.) in Coastal and Non-coastal Stands

Wind damage to the leaves of sycamore (Acer pseudoplatanus L.)trees was examined at five sites in Northern Ireland duringJune and September, 1986 by estimating the percentage of leafarea showing lesions, discoloration or actual loss of laminasurface. Levels of damage on individual leaves ranged up to 46.6 percent of the leaf area but few leaves had damage greater than10 percent. Two sites showed relatively low levels of damage(about 2 per cent of lamina damaged); one site was inland, andthe other, although coastal, was well sheltered. The three othersites had damage in excess of 3.5 per cent of the lamina area—thesite with the highest levels of damage being the most exposedsite and the one probably experiencing the greatest amount ofturbulence. Most damage had occurred by June and there was littleincrease in damage levels between the June and September samples;there was however evidence of the premature loss of the mosthighly damaged leaves. Regression analysis indicated that at sites showing the highestlevels of damage, the most highly damaged leaves were the smallest.It is argued that in these cases the damage to the leaves issufficiently high to prevent full leaf expansion. At sites wherethe overall damage levels were lower, there was either a non-significantrelationship between percentage leaf damage and leaf area ora significant positive relationship. It is suggested here thatthe damage levels are not great enough to inhibit the expandingleaves reaching full size.     

Seed Size as a Predictor of Germination Success and Early Seedling Growth in ‘Hollong’ (Dipterocarpus macrocarpus Vesque)

In species with seeds losing viability shortly after dispersal and exhibiting inherently low germination, quick decisions are required with respect to seeds that should be selected to maximize germination success and vigorous growth of seedlings. In ‘hollong’ (Dipterocarpus macrocarpus Vesque), I address the following questions: (a) are seeds that germinate randomly distributed within a seed population, (b) are subpopulations of viable and non-viable seeds separable, (c) does seed size predict which seeds germinate and (d) does seed size predict time required for germination and seedling vigour? Two estimators of seed size, diameter and weight, demonstrated a significant positive linear relationship endorsing assumption that accumulation of mass increases with increase in diameter of seeds. A threshold for selection of potential seeds for germination could not be derived from diameter measurements since seeds in an entire range of diameter did or did not germinate. All seeds <11 g did not germinate, but this threshold lies at the far lower end of the weight range and allows rejection of only a few seeds and acceptance of many seeds that will not germinate. A risk of potential seeds being rejected or non-viable seeds being picked exists if selection was derived from either diameter or weight. However, viable seeds could be better predicted from a scatter-plot of diameter on x-axis and weight on y-axis. Seeds showed a fan-shaped scatter and those developing the lower blade of the fan did not germinate while those following the handle were successful. Hence, two subpopulations segregated, but with a fuzzy edge. Mean diameter and mean weight of germinated seeds were significantly greater than those of ungerminated seeds. Most ungerminated seeds were those that had relatively smaller weight compared to their diameter. Although some large diameter seeds with small weight did germinate, but failed to develop into seedlings. Germination time and seedling vigour parameters (height, leaf number and collar diameter) were correlated both with diameter and weight. However, weight appears to have mattered more than diameter in germination success and early seedling growth.     

A low <Emphasis Type="Italic">Platypus quercivorus</Emphasis> hole density does not necessarily indicate a small flying population

The ambrosia beetle Platypus quercivorus causes mass mortality of Fagaceae trees in Japan, and tree species differ in their susceptibility to P. quercivorus. We hypothesized that interspecific differences in susceptibility are caused by differences in beetle infestation patterns, that is, how many beetles fly to a tree and how many of those arriving bore holes. To examine how tree susceptibility is related to these parameters, two tree species with different degrees of susceptibility were studied (highly susceptible Quercus crispula and less susceptible Q. salicina). Specifically, we measured the number of male beetles per unit area that flew to the host trees (NFM) and the density of holes bored by male beetles (DH). From these two values, we calculated the proportion of male beetles flying to a host tree that bored holes (PBM). These parameters were compared for the two Quercus species. Although the two species did not differ in NFM in 2003, PBM was markedly lower for Q. salicina than for Q. crispula and DH was significantly lower for Q. salicina than for Q. crispula. Thus, the lower susceptibility of Q. salicina is partly explained by its low PBM.