Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

Impact of silviculture on dead wood and on the distribution and frequency of tree microhabitats in montane beech-fir forests of the Pyrenees

In forest ecosystems, the level of biodiversity is strongly linked to dead wood and tree microhabitats. To evaluate the influence of current forest management on the availability of dead wood and on the abundance and distribution of microhabitats, we studied the volume and diversity of dead wood objects and the distribution and frequency of cavities, dendrothelms, cracks, bark losses and sporophores of saproxylic fungi in montane beech-fir stands. We compared stands unmanaged for 50 or 100 years with continuously managed stands. A total of 1,204 live trees and 460 dead wood objects were observed. Total dead wood volume, snag volume and microhabitat diversity were lower in the managed stands, but the total number of microhabitats per ha was not significantly different between managed and unmanaged stands. Cavities were always the most frequent microhabitat and cracks the least frequent. Dendrothelm and bark loss were favored by management. Beech (Fagus sylvatica) carried many more microhabitats than silver fir (Abies alba), especially cavities, dendrothelms and bark losses. Fir very scarcely formed dendrothelms. Secondary tree species played an important role by providing cracks and bark losses. The proportion of microhabitat-bearing trees increased dramatically above circumference thresholds of 225 cm for beech and 215 cm for fir. Firs with a circumference of less than 135 cm did not carry microhabitats. In order to conserve microhabitat-providing trees and to increase the volume of dead wood in managed stands, we recommend conserving trees that finish their natural cycle over 10–20% of the surface area.     

A study on Xingkai Lake pine gall rust

The Xingkai Lake pine (Pinus takahasii Nakai) gall rust caused byCronartium quercuum(Berk.) Miyabc: Shirai is a serious stem rust in the northeast region of China. The alternate host is oak (Quercus mongolica Fisch). Germination of both acciospores and urediospores was optimal at 12°C and occurred over a range of temperatures, from 4°C to 32°C. Teliospores germinated optimally at 16–18°C and over a range of 8–28°C. All spores germinated best under natural light and dark conditions. Direct light inhibited germination even when followed by darkness. The incubation periods of urediospores and teliospores in inoculation experiments were 6–16 days and 3–33 days respectively. Anatomical studies ofCronartium quercuum galls showed that rust hyphae freely ramify through the intercellular spaces within parenchymatous tissues of the cortex, phloem, cambium and xylem. Hyphae are perennial; large haustoria arc cylindrical with rounded or blunt ends. Among the 12 fungicides tested for control of the rust, pine-tar, pine-tar: diesel oil (1:1, 1:3, 1:5), 75% Bravo emulsion and 70% Mancozeb emulsion applied to the galls produced satisfactory results.     

丝栗栲林生长与生产力的研究

在福建省邵武市洪墩采育场，调查丝栗栲林标地１０块，做平均木、树干解析木９株，杉木人工林标地２块，研究丝栗栲的生长与生产力，结果表明：丝栗栲林的胸径、树高及单株材积总生长量，均随年龄的增加而增加；胸径２０ａ前生长最快；树高１６ａ前生长最快；材积３６ａ时尚未有下降的趋势。胸径、树高平均生长量，随年龄增加而增加，当生长量达高峰后，则随年龄的增加而下降；胸径生长高峰，好立地１２ａ，中、劣立地１５ａ；树高生长高峰，好立地１０ａ，中立地８ａ，劣立地１２ａ；材积平均生长量，随年龄增加而增加，好、劣立地３６ａ时仍未下降，中立地３４ａ时生长量最大。胸径、树高连年生长量随年龄增加而下降；材积连年生长量，随年龄增加而增加，当生长量达高峰后，则随年龄增加而下降，生长高峰出现的年龄，好立地２８ａ，中立地２２ａ，劣立地２６ａ。在相似的立地条件下，丝栗栲林的蓄积量比杉木人工林低，但乔木层总生物量比杉木林高。     

Influence of different forest conversion strategies on ground vegetation and tree regeneration in pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) stands: a case study in NE Germany

The conversion of anthropogenic into more natural, self-regenerating forests is one of the major objectives of forestry throughout Europe. In this study, we present investigations on permanent plots with different silvicultural treatment in NE German pine stands. Management of old-growth pine stands on acidic and nutrient-poor sandy sites differs in fencing, thinning, and planting of certain tree species. The investigations were carried out on the community, population, and individual level of the pine forest ecosystems. Thus, vegetation changes, size and height of tree populations, and height increment of tree individuals were observed over a time span of 6 years. Special attention was paid to short-lived tree species such as, e.g., Frangula alnus and Sorbus aucuparia, as well as to Fagus sylvatica as one of the most typical forest tree species of Central Europe. Vegetation changes are interpreted as a consequence of natural regeneration of formerly degraded forest sites, involving an increase in nutrient availability. High browsing pressure can be considered as a key factor for the inhibition of tree seedlings and growth of saplings. Some Sorbus aucuparia individuals, however, succeeded in growing out of the browsing height also in unfenced stands. Few found specimens of Fagus sylvatica proved that this species is able to establish spontaneously on these relatively dry, acidic sites under continental climate influence. Such natural regeneration processes, also including spontaneous rejuvenation of trees, can be integrated into silviculture as passive forest conversion management. An active management like thinning of stands, planting of trees, and fencing can accelerate forest conversion with regard to height growth and species number of trees.     

Long-term changes of structure and tree species composition in Dinaric uneven-aged forests: are red deer an important factor?

Uneven-aged silver fir-European beech forest stands were studied to (1) analyse the dynamics of diameter structure and tree species composition in the past two centuries and (2) determine the impact of red deer on the regeneration and recruitment of silver fir. The study used current data on forest stands, archival data from old forest management plans for the period 1789–2004, and red deer harvesting records for the period 1907–2006. During the observation period, the silver fir population aged and silver fir and European beech alternated in dominance. The study revealed a strong impact of red deer on the composition and recruitment of tree regeneration, especially on silver fir regeneration. The drastic changes in red deer density (from extermination up to 5.8 animals km⁻²) and past forest management practices were apparently the main factors driving the population dynamics of silver fir (regeneration, recruitment, and diameter structure) in the study area during the past two centuries.     

Dendroclimatological regions of Douglas fir (<Emphasis Type="Italic">Pseudotsuga menziesii</Emphasis> Franco) in western Poland

Abstract Six Douglas fir (Pseudotsuga menziesii Franco) stands were selected in both the Sudety Mountains and the Great Poland Lowland. These two regions are distinctly different with regard to thermal and pluvial conditions. In each stand, two increment cores per tree were extracted from 20 approximately 100-year-old trees and the tree-ring widths measured. It is clear from the different growth reactions of the study trees to the climatic conditions that the Sudety Mountains and the Great Poland Lowland are two dendrochronologically separate regions. The tree-ring width chronologies form homogeneous groups for each region. The tree ring–climate models for Douglas fir in both regions indicate similarities and differences. The similarities among chronologies are attributed to winter temperature. The differences are attributed to rainfall (which changes with longitude and altitude of sites) during the vegetation season.     

Effects of thinning on growth of six tree species in north-temperate forests of Lithuania

This paper summarises the results from 35 years-observed thinning experiments on 256 permanent sample plots in 10–60 year-old stands of ash, aspen, birch, oak, pine and spruce in Lithuania. Thinning enhanced crown projection area increment of residual trees. The largest effect was observed in stands of aspen and birch (growth increase by 200%), followed by ash and oak (over 100%), and spruce and pine (about 80%). Thinning also promoted dbh increment, especially in younger stands, and the increase of dbh increment was positively correlated with the thinning intensity. The strongest reaction was exhibited by oak and aspen, while ash, birch and conifers reacted to a lower extent. Low and moderate intensities of thinning stimulated volume production in younger stands while the opposite was observed in older stands with increasing removals. Spruce stands exhibited relatively strongest increase of volume increment and pine, –the weakest, while the effect on deciduous species was intermediate. The results demonstrate that significant increase in volume increment is achievable with thinning of only young forest stands, e.g. 10–20 year-old pine, birch and ash, or 10–30 year-old oak, aspen and spruce.     

Predicting tree regeneration in <Emphasis Type="Italic">Picea abies</Emphasis> snag stands

A bark beetle (Ips typographus) infestation caused the death of almost all Norway spruce (Picea abies) trees in a mountain forest in the Swiss Alps. We developed a tree regeneration model, ‘RegSnag’ (=REGeneration in a SNAG stand), to project the future amount and height of tree regeneration in these snag stands. The model combines a height-class structured tree module with a microsite-based module of snag decay and ground-vegetation succession. Microsite-specific rates of germination, mortality and height growth were modelled for four tree species (Picea abies, Sorbus aucuparia, Acer pseudoplatanus and Betula pendula) in eight height classes (from seedlings to saplings 5 m tall) and on 26 microsite types (e.g. moss, grass). Model tests with independent field data from 8 years after the Picea die-back demonstrated that microsites had a considerable effect on the development of tree regeneration on both the montane and the subalpine level. With microsite-specific parameters, the height and frequency of Picea in each microsite could be simulated more accurately than without considering microsite effects (e.g. bias of 8 vs. 119 saplings ha⁻¹ on the montane level). Results of simulations 40 years into the future suggest that about 330–930 Picea saplings per ha out of those that germinated in 1994 and 1996 will reach a height of 5 m within 30–35 years after Picea die-back. This is due to differences in seed inflow and browsing intensities. Picea and not Betula or Sorbus trees will replace the current herbaceous vegetation in these snag stands.     

European pear rust on Juniperus excelsa L. in south‐western Turkey

In this study, the frequency of European pear rust (Gymnosporangium fuscum) lesions on Juniperus excelsa was investigated both on seedlings in a nursery and on forest trees in six juniper stands located in south‐western Turkey. In the nursery, lesions occurred on 0.7% of 5510 1‐year‐old seedlings and 29% of 200 2‐year‐old potted seedlings. In each of the juniper forests, 50 trees were chosen within a square of approximately 400 m² in size. In general, rust lesions were common on the trees. On average, 49.0% of the trees had lesions; on 17.3% of the trees, the lesions were located on the trunk, on 43.3% on the branches and on 11.7% on both. In lesion‐bearing trees, the average number of lesions was 1.47–3.59 and the maximum number of lesions per tree were six to 11. Most of the lesions (85.1%) were on branches. There were significant differences between the stands, which can be attributed to different age of the trees, site factors and the presence of alternate hosts. The rust identified from stem lesions in forest trees and from nursery seedlings was European pear rust (G. fuscum), but possible co‐occurrence of other Gymnosporangium spp. cannot be excluded.     

两种接种方式下角倍蚜虫瘿对寄主盐肤木枝叶生长的影响

[目的]明确两种接种方式(自然迁飞和人工挂袋)下角倍蚜虫瘿的数量和分布及其对寄主盐肤木枝叶生长的影响。[方法]在自然迁飞和人工挂袋形成虫瘿的盐肤木林内,按随机区组设计设置样地,在样地内随机选取有虫瘿树和无虫瘿树,分别测量和统计盐肤木及虫瘿的生长指标,采用方差分析虫瘿对盐肤木生长的影响。[结果]在自然迁飞和人工挂袋两种接种方式下,盐肤木单株虫瘿数没有显著差异。但人工挂袋时,单片复叶的平均虫瘿总体积为170.92±14.85 cm~3、小叶总面积为616.26±32.73 cm~2,显著高于自然迁飞形成虫瘿的总体积85.82±9.40cm~3和小叶总面积482.81±28.51 cm~2。表明人工挂袋接种可以增加虫瘿的总体积,从而提高角倍产量;同时,较大体积的虫瘿还促进了盐肤木小叶的生长,使小叶面积增大。[结论]角倍蚜虫瘿对寄主盐肤木的枝叶生长具有促进作用,且这种生长促进与虫瘿的体积呈正相关。与自然迁飞接种相比,人工挂袋接种更能发挥盐肤木的生长和结倍潜力,提高角倍产量。     

Light distribution in scattered-trees open woodlands in Western Spain

We have studied the percentage of radiation transmitted through the tree canopy to the underlying pasture and crops in dehesas of Southwest Spain by means of fish-eye photographs taken at different distances from the tree. Thirty-six trees were studied covering all the diametric classes (0.1–14 m canopy width) of two stands, with mean density of 19 mature trees ha⁻¹. Intercepted light decreased with distance following an logistic curve, indicating a rapid increase in the light availability with distance from the tree. For mature trees, radiation was constant beyond 20 m. Applying a multivariable regression light equation, distance, stem diameter and canopy width explained more than 88% of the light variability for each orientation studied. A simple model was built up from light equations, tree growth curves and allometric relationships. From this model, we have estimated that radiation available for crops and pasture decreased up to 21% due to the presence of trees in a standard dehesa with 24 mature trees ha⁻¹ and 13% of canopy cover. In addition, we have generated different radiation maps of virtual dehesas differing in tree age, density and arrangement, which could be useful to determine optimal tree planting schemes and consequent pruning and thinning regimes.     

Relationships between mass mortality of two oak species (Quercus mongolica Turcz. var.grosseserrata Rehd. et Wils. andQ. serrata Thunb.) and infestation by and reproduction ofPlatypus quercivorus (Murayama) (coleoptera: Platypodidae)

This study investigated the number of adults ofPlatypus quercivorus (Murayama) captured on host oak (Quercus spp.) trees, the attack density (the number of entry holes per 100 cm²), and the number of emerging adults to clarify the relationships between the beetle’s colonization on trees in oak stands and tree mortality. The initial attack ofP. quercivorus, which was the most intense attack, was observed on most living trees simultaneously. Although some attacked trees died within the year of the attack or in the next year, a high density of attack did not usually cause the death of host trees. Surviving trees suffered low levels of consecutive attack after the initial attack. BecauseP. quercivorus successfully produced broods only in the dead trees, the population of the insect seems to be maintained only in stands where oak mortality occurs. However, the adults that landed on most of the surviving trees appeared unable to reproduce probably due to degradation of host quality. Thus, oak mortality probably ceases within 3 or 4 years after the start of infestation in a stand, with subsequent reductions in population density of the borer.     

Response of slash pine planted on mounds in central and southwestern Louisiana

On two silt loam flatwoods in Louisiana, use of mounds as planting sites for slash pine (Pinus elliottii Engelm. var. elliottii) was shown to be more effective than planting without mounds (untreated control) after six growing seasons in the field. Mounds formed were of two sizes: low mounds made of 550 dm³ of soil and high mounds made of 1,000 dm³ of soil. One site was in central Louisiana and had two drainage classes: a somewhat poorly drained Caddo-Messer complex and a poorly drained Caddo soil. On both drainage classes, pine survival was not affected by treatment, but the incidence of fusiform rust galls on the main stem (caused by Cronartium quercuum (Berk.) Miyabe ex Shirai f. sp. fusiforme Burdsall and Snow) was greater on trees planted on mounds. The second site was a poorly drained Caddo soil in southwestern Louisiana. At this site, there were fewer surviving pines on controls than on mounds, but the incidence of rust galls was not affected by treatment. At both sites, pines were taller and had a greater mean diameter at breast height on mounds than on controls. Mound size did not influence tree size at either site.     

Individual-based measurement and analysis of root system development: case studies for <Emphasis Type="Italic">Larix gmelinii</Emphasis> trees growing on the permafrost region in Siberia

We present results of individual-based root system measurement and analysis applied for Larix gmelinii trees growing on the continuous permafrost region of central Siberia. The data of root excavation taken from the three stands were used for the analyses; young (26 years old), mature (105 years old), and uneven-aged over-mature stand (220 years old). In this article, we highlight two topics: (1) factors affecting spatio-temporal pattern of root system development, and (2) interactions between aboveground (i.e., crown) and belowground (i.e., root) competition. For the first topic, the detailed observation of lateral roots was applied to one sample tree of the overmature stand. The tree constructed a superficial (<30 cm in depth) and rather asymmetric root system, and each lateral root expanded mainly into elevated mounds rather than depressed troughs. This indicated that spatial development of an individual root system was largely affected by microtopography (i.e., earth hummocks). For these lateral roots, elongation growth curves were reconstructed using annual-ring data, and annual growth rates and patterns were compared among them. The comparison suggested that temporal root system development is associated with differences in carbon allocation among the lateral roots. For the second topic, we examined relationships between individual crown projection area (CA) and horizontal rooting area (RA) for the sample trees of each stand. RA was almost equal to CA in the young stand, while RA was much larger (three or four times) than CA in the mature and overmature stands. Two measures of stand-level space occupation, crown area index (aboveground: CAI; sum of CAs per unit land area) and rooting area index (belowground: RAI; sum of RAs), were estimated in each stand. The estimates of RAI (1.3–1.8 m² m₋₂) exceeded unity in all stands. In contrast, CAI exceeded unity (1.3 m² m₋₂) only in the young stand, and was much smaller (<0.3 m² m₋₂) in the two older stands. These between-stand differences in RAI–CAI relationships suggest that intertree competition for both aboveground and belowground spaces occurred in the young stand, but only belowground competition still occurred in the two older stands. Based on this finding, we hypothesized that competition below the ground may become predominant as a stand ages in L. gmelinii forests. Methodological limitations of our analysis are also discussed, especially for the analysis using the two indices of space occupation (CAI, RAI).     

Comparison of selected statistical distributions for modelling the diameter distributions in near-natural <Emphasis Type="Italic">Abies</Emphasis>–<Emphasis Type="Italic">Fagus</Emphasis> forests in the Świętokrzyski National Park (Poland)

The aim of the study is to compare selected theoretical distributions (normal, lognormal, Weibull, gamma, logistic, and exponential) in describing the tree diameter (DBH) distributions of mixed near-natural forests consisting of fir Abies alba Mill. and beech Fagus sylvatica L. growing in various vertical structures. Tree DBH data were collected between 1997 and 2008 from 51 sample plots established in the Świętokrzyski National Park in Poland. The empirical data represent differentiated DBH distributions, ranging from almost symmetric to extremely asymmetric ones. The chi-square test and the modified Kolmogorov–Smirnov test were chosen for the goodness-of-fit testing. In addition to the test statistics, the bias (B), the root mean square error (RMSE) and the graphical method (quantile–quantile plots) were used. In one-storied stands, the most suitable distributions were the normal and logistic distributions; in two-storied and multilayered stands, the Weibull and gamma distribution were the most suitable; and in selection stands, the exponential distribution was the most appropriate to describe the DBH distribution. The order of precision of the tested distributions (from the highest to the lowest) was Weibull, gamma, logistic, normal, exponential, and lognormal. The normal and exponential distribution should be applied only to one-storied and selection forests, respectively. The least suitable distribution for DBH distribution modelling was the lognormal one.     

Tiarosporella durmitorensis sp. nov. – a new pathogenic fungus on needles of Abies alba

A fungus causing necrosis and needle cast has been discovered on fir trees (Abies alba) in a hight of 1700 m above sea level in Yugoslavia. The intensity of infection was very high, and the needles of all age classes were infected. The consequence of the infection is the death of the tree. The pathogen is described as a new species Tiarosporella durmitorensis Karad?i?. So far, this fungus has been found only in European silver fir stands in the National Park Durmitor and in the National Park Biogradska Gora (Montenegro).     

Above- and belowground biomass,nutrient and carbon stocks contrasting an open-grown and a shaded coffee plantation

Coffee (Coffea canephora var robusta) is grown in Southwestern Togo under shade of native Albizia adianthifolia as a low input cropping system. However, there is no information on carbon and nutrient cycling in these shaded coffee systems. Hence, a study was conducted in a mature coffee plantation in Southwestern Togo to determine carbon and nutrient stocks in shaded versus open-grown coffee systems. Biomass of Albizia trees was predicted by allometry, whereas biomass of coffee bushes was estimated through destructive sampling. Above- and belowground biomass estimates were respectively, 140 Mg ha⁻¹ and 32 Mg ha⁻¹ in the coffee–Albizia association, and 29.7 Mg ha⁻¹ and 18.7 Mg ha⁻¹ in the open-grown system. Albizia trees contributed 87% of total aboveground biomass and 55% of total root biomass in the shaded coffee system. Individual coffee bushes consistently had higher biomass in the open-grown than in the shaded coffee system. Total C stock was 81 Mg ha⁻¹ in the shaded coffee system and only 22.9 Mg ha⁻¹ for coffee grown in the open. Apart from P and Mg, considerable amounts of major nutrients were stored in the shade tree biomass in non-easily recyclable fractions. Plant tissues in the shaded coffee system had higher N concentration, suggesting possible N fixation. Given the potential for competition between the shade trees and coffee for nutrients, particularly in low soil fertility conditions, it is suggested that the shade trees be periodically pruned in order to increase organic matter addition and nutrient return to the soil. An erratum to this article can be found at     

Species diversity of broad-leaved trees in Cryptomeria japonica plantations in relation to the distance from adjacent broad-leaved forests

The species diversity of broad-leaved trees in relation to the distance from a broad-leaved forest was investigated in Cryptomeria japonica plantations with crown snow damage near the Japan Sea in central Japan. The number and diversity of species and stem density decreased with distance from the broad-leaved forest in maturing gap stands (51–58 years old; >10 years after crown snow damage), but not in recent gap stands (34–42 years old; <10 years after crown snow damage), and increased with improved light conditions in the interior of recent gap stands, but not in that of maturing gap stands. The stem densities of tall and small tree species and woody lianas were greater in the interior of recent gap stands than in maturing gap stands. Woody lianas, which are characteristically shade intolerant, had a high stem density in the interior of recent gap stands. In contrast, the stem density of shrubs was greater in maturing gap stands than in recent gap stands, irrespective of distance. Shade-tolerant shrubs had a high stem density in maturing gap stands. Wind-dispersed and frugivore-dispersed species were concentrated at the edges of maturing gap stands, but some frugivore-dispersed species, which may persist in soil seed banks, occurred in the interior of recent gap stands. Gravity-dispersed species tended to occur both at the plantation edge and in the interior. The differences in the occurrence patterns of broad-leaved trees in the two types of stands reflected the difference in the effect of both the distance from the adjacent broad-leaved forest and the light conditions related to canopy gaps, with the time since gap formation in a region where crown snow damage often occurs.     

Effects of different ecological and silvicultural factors on beetle catches in the Turkish fir (<Emphasis Type="Italic">Abies bornmülleriana</Emphasis> Mattf.) ecosystems

To study the bark beetle attacks on Turkish fir stands and to determine the principle site and stand factors influencing beetle attacks, trap log method was used in the Western Black Sea Region of Turkey. The study was carried out in ten different locations prior to flying time in February 2003. On each experimental plot, three healthy fir trees with 30–50 cm diameter and 200 m apart were cut down with a chainsaw. From each sampling tree, needles were sub-sampled for determination of specific leaf-area and macro-nutrient concentrations. In addition, on each experimental plot, crown closure, basal-area, and stand density were measured. To determine the bulk density and nutrient concentrations of the soil, samples were taken around trees on each plot. The relationship between measured stand variables, the number of beetle species and beetle populations were evaluated using correlation and stepwise multiple regression procedure. Eight different species belonging to four different families from Coleoptera were identified in examining the trap logs. Six of these eight species (Pissodes piceae Illig., Pissodes notatus Fabr., Cryphalus piceae Ratz., Pityophthorus pityographus Ratz., Xyloterus lineatus Oliv., Pityokteines curvidens Germ.) were identified as harmful beetles (HB). The other two (Thanasimus formicarius L. and Rhizophagus dispar Payk.) were identified as predator species. The analysis showed that the total number of HB increased as the site’s slope and needle specific leaf area increased. However, as needle N concentrations increased, HB population decreased. Increased Ca concentration in fir needle reduced total insect attack.