Area‐wide application of verbenone‐releasing flakes reduces mortality of whitebark pine Pinus albicaulis caused by the mountain pine beetle Dendroctonus ponderosae

• 1 DISRUPT Micro‐Flake Verbenone Bark Beetle Anti‐Aggregant flakes (Hercon Environmental, Inc., Emigsville, Pennsylvania) were applied in two large‐scale tests to assess their efficacy for protecting whitebark pine Pinus albicaulis Engelm. from attack by mountain pine beetle Dendroctonus ponderosae Hopkins (Coleoptera: Scolytinae) (MPB). At two locations, five plots of equivalent size and stand structure served as untreated controls. All plots had early‐ to mid‐outbreak beetle populations (i.e. 7.1–29.2 attacked trees/ha). Verbenone was applied at 370 g/ha in both studies. Intercept traps baited with MPB aggregation pheromone were placed near the corners of each plot after the treatment in order to monitor beetle flight within the plots. Trap catches were collected at 7‐ to 14‐day intervals, and assessments were made at the end of the season of stand structure, stand composition and MPB attack rate for the current and previous years.
• 2 Applications of verbenone flakes significantly reduced the numbers of beetles trapped in treated plots compared with controls at both sites by approximately 50% at the first collection date.
• 3 The applications also significantly reduced the proportion of trees attacked in both Wyoming and Washington using the proportion of trees attacked the previous year as a covariate in the model for analysis of current year attack rates; in both sites, the reduction was ≥ 50%.
• 4 The flake formulation of verbenone appears to have promise for area‐wide treatment by aerial application when aiming to control the mountain pine beetle in whitebark pine forests.

     

Attack by Hylobius warreni on grafted lodgepole pine and its relationships with monoterpene composition and scion : rootstock diameter ratio

1 A lodgepole pine seed orchard at the Prince George Tree Improvement Station (PGTIS), with up to 60% of grafted trees attacked by the Warren root collar weevil Hylobius warreni was investigated to determine whether relative monoterpene composition or scion : rootstock interactions of grafts affected susceptibility to attack. 2 There was a significant relationship between relative levels of α‐pinene, β‐thujene, β‐pinene, δ‐3‐carene and limonene in scion and rootstock in unattacked trees, indicating a potential effect of the scion monoterpene composition on their composition in the rootstock. 3 Relative content of δ‐3‐carene and β‐phellandrene differed significantly in root stocks of attacked and unattacked trees but, for individual clones, a significant difference was only detected for β‐phellandrene in one clone. δ‐3‐Carene levels may have been too low in the examined trees to exert a strong effect. 4 Interestingly, attack status was significantly associated with two scion monoterpenes: α‐thujene and α‐terpinolene, both of which had higher levels in unattacked than in attacked trees. 5 Warren root collar weevils appear largely unaffected by monoterpene content, but further study is required to determine whether high levels of δ‐3‐carene imparts some level of resistance to attack, and to verify whether the observed effects of scion monoterpenes are real or artefacts of the analysis. 6 Hylobius warreni‐attacked trees had smaller scion : rootstock diameter ratio (i.e. a large rootstock diameter relative to the scion diameter) than unattacked trees. This effect was consistent among clones, and was not due to the absolute diameter of the rootstock or the scion. Trees with increased diameter‐growth at the root collar (e.g. some grafted trees) may have increased susceptibility to attack by H. warreni, or diameter‐growth at the root collar is affected by the attack.     

Pre-dispersal seed losses to insects in species of Leptospermum (Myrtaceae)

Pre-dispersal seed losses to insects in species of Leptospermum at Wilson's Promontory in south-eastern Australia varied markedly between species, seasons, sites and individuals. The proportion of fruits attacked by insects averaged 11% (range 3–37%) in populations of Leptospermum juniperinum, 23% (6–95%) in Leptospermum myrsinoides, 23% (0–47%) in Leptospermum laevigatum, and 28% (8–45%) in Leptospermum lanigerum. The proportion of seeds lost in attacked fruits also varied widely. Differences between species varied markedly from year to year, but differences between sites were relatively constant. Fruit production by L. myrsinoides was unusually low at most sites during 1984; in each case this coincided with unusually high rates of insect attack, suggesting that predator satiation operates during normal seasons.     

Site condition and predation influence a bark beetle's success: a spatially realistic approach

Abstract 1 Spatial pattern in abundance of Dendroctonus micans was studied in a 600‐ha spruce stand in the Massif Central (Lozère, France). The proportion of trees attacked was measured in 38 plots and these data were used to estimate spatial pattern of attack density in the stand and to identify a transect of decreasing attack density (80% to 30%) over less than 1000 m. 2 Spatial variation in attack density was analysed in relation to (i) data on site and stand characteristics (altitude, slope, tree density, tree average height, yield class and average age) collected from 63 points in the stand and (ii) the releases of the predator Rhizophagus grandis (localization and number of beetles released). 3 The proportion of attacked trees was analysed using geostatistics and showed a strong spatial structure reflecting the spatial scale of interaction of D. micans with its environment. The spatial structure was modelled in order to estimate the spatial distribution of attack density at unsampled locations. 4 A linear model relating interpolated attack density to the number of predators released 6–10 years before the survey in a 300‐m radius and to the average slope over a 250‐m radius explained 67% of the observed variability. Spatial autocorrelation was taken into account in a spatial regression model.     

Characteristics of Norway spruce trees (Picea abies) surviving a spruce bark beetle (Ips typographus L.) outbreak

The characteristics of spruce individuals, which survived a massive bark beetle outbreak, were compared with the characteristics of neighbouring attacked trees in Šumava National Park (Czech Republic). Selected parameters related to crown geometry, stand conditions and distances between trees were measured or estimated. Significant differences were found between the surviving trees and the neighbouring trees attacked by I. typographus. Trees with a higher level of stem shading (longer crown length) tended to survive. The attacked trees were usually located in areas with larger basal area, especially southwards from them. A shorter distance to a previously attacked tree increased the probability of additional attack. Spruce trees with more progressive crown structure transformation (primary structure defoliation) were significantly more frequently attacked by spruce bark beetle. Superior and taller trees had a clearly longer life expectancy than dominant ones. These results show that the attack of trees by bark beetle can be predicted to a certain degree, which can be used in management of endangered spruce forests.     

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.     

The effect of stem density on the probability of attack by the ambrosia beetle Platypus quercivorus varies with spatial scale

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Allometric Models for Predicting Aboveground Biomass in Two Widespread Woody Plants in Hawaii

Allometric models are important for quantifying biomass and carbon storage in terrestrial ecosystems. Generalized allometry exists for tropical trees, but species‐ and site‐specific models are more accurate. We developed species‐specific models to predict aboveground biomass in two of the most ubiquitous natives in Hawaiian forests and shrublands, Metrosideros polymorpha and Dodonaea viscosa. The utility of the M. polymorpha allometry for predicting biomass across a range of sites was explored by comparing size structure (diameter at breast height vs. tree height) of the trees used to develop the models against trees from four M. polymorpha‐dominated forests along a precipitation gradient (1630–2380 mm). We also compared individual tree biomass estimated with the M. polymorpha model against existing generalized equations, and the D. viscosa model with an existing species‐specific model. Our models were highly significant and displayed minimal bias. Metrosideros polymorpha size structures from the three highest precipitation sites fell well within the 95% confidence intervals for the harvested trees, indicating that the models are applicable at these sites. However, size structure in the area with the lowest precipitation differed from those in the higher rainfall sites, emphasizing that care should be taken in applying the models too widely. Existing generalized allometry differed from the M. polymorpha model by up to 88 percent, particularly at the extremes of the data range examined, underestimating biomass in small trees and overestimating in large trees. The existing D. viscosa model underestimated biomass across all sizes by a mean of 43 percent compared to our model. The species‐specific models presented here should enable more accurate estimates of biomass and carbon sequestration in Hawaiian forests and shrublands.     

Winter starch reserves of white oak as a predictor of attack by the twolined chestnut borer,Agrilus bilineatus (Weber) (Coleoptera: Buprestidae)

Summary The twolined chestnut borer, Agrilus bilineatus (Weber) (Coleoptera: Buprestidae), attacks oaks (Quercus spp.) and is associated with extensive mortality of trees in the eastern deciduous forests of North America. We tested the hypothesis that winter starch reserves of oak roots are an indicator of tree vigor and that only trees low in stored starch would be attacked by A. bilineatus. We measured the levels of stored starch in the roots of 200 non-infested healthy white oaks during the dormant season and determined their correlation with A. bilineatus attacks the following spring. There was a significant increase in A. bilineatus captures on sticky traps with a decrease in winter starch reserves. Trees low in stored starch that were also stressed by phloem-girdling attracted 3.7 times as many beetles as did non-girdled trees that were low in starch. However, non-girdled trees that had low winter starch reserves were also attacked. Only oaks that had had extremely low winter root starch reserves (<5mg/g dry weight of root sapwood tissue) were heavily attacked by A. bilineatus and subsequently died. One third of non-girdled low starch trees and 67% of phloem-girdled low starch trees died, whereas none of the trees with root starch >5 mg/g dry wt died. These results indicate that winter starch reserves are a good predictor of A. bilineatus attack.The investigation reported in this paper (No. 87-7-8-118) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

Attraction of ambrosia and bark beetles to coast live oaks infected by Phytophthora ramorum

1 Sudden oak death is caused by the apparently introduced oomycete, Phytophthora ramorum. We investigated the role of bark and ambrosia beetles in disease progression in coast live oaks Quercus agrifolia. 2 In two Marin County, California sites, 80 trees were inoculated in July 2002 with P. ramorum and 40 were wounded without inoculation. Half of the trees in each group were sprayed with the insecticide permethrin [cyclopropanecarboxylic acid, 3‐(2,2‐dichloroethenyl)‐2,2‐dimethyl‐(3‐phenoxyphenyl) methyl ester] to prevent ambrosia and bark beetle attacks, and then were sprayed twice per year thereafter. After each treatment, sticky traps were placed on only the permethrin‐treated trees. Beetles were collected periodically in 2003. 3 Inoculated trees accounted for 95% of all beetles trapped. The ambrosia beetles Monarthrum scutellare and Xyleborinus saxeseni and the western oak bark beetle Pseudopityophthorus pubipennis were the most abundant of the seven species trapped. 4 Permethrin treatment delayed initiation of beetle attacks and significantly reduced the mean number of attacks per tree. Beetles did not attack any wounded or noncankered inoculated trees. 5 Trees with larger cankers trapped more beetles early in the disease. Once permethrin lost effectiveness, the number of beetle entrance tunnels was a more reliable predictor of subsequent trap catch than was canker size. 6 Beetles were initially attracted to P. ramorum cankers in response to kairomones generated in the host‐pathogen interaction. After beetles attacked the permethrin‐treated trees, aggregation pheromones most probably were the principal factor in beetle colonization behaviour.     

The persistence of blue ash in the aftermath of emerald ash borer may be due to adult oviposition preferences and reduced larval performance

1. We examined the health of mature blue ash, Fraxinus quadrangulata, in two forests in southwestern Ohio in relation to that of mature white ash, F. americana, and examined the potential importance of oviposition preferences and larval resistance in the persistence of blue ash.
2. Both blue ash and white ash were largely unaffected by emerald ash borer in 2012. By 2018, nearly 90% of the blue ash trees observed in these forests had full or nearly full canopies, as opposed to less than 20% of the white ash encountered in our studies. In 2021, blue ash maintained a similar degree of canopy health as in 2018, but no standing live mature white ashes remained.
3. Bark removals revealed no current or past larval feeding attempts in blue ash in 2018 or 2021, except for one attacked and killed tree in 2018. All white ash trees examined were attacked. In a laboratory bioassay with cut stems, emerald ash borer larvae fed less and grew significantly more slowly on blue ash than on white ash.
4. Both reduced larval performance and reduced adult oviposition likely contribute to the persistence of blue ash in forests devastated by emerald ash borer.

     

Why does the bud-galling sawfly,Euura mucronata,attack long shoots?

Summary The bud-galling sawfly, Euura mucronata, attacked longer shoot length classes on its host, Salix cinerea, more frequently than shorter shoots. Shoot length accounted for 76 to 93 percent of the variance in number of galls per 100 shoots in three habitats: forest, watermeadow, and lakeside. The reasons for this pattern were addressed with studies on shoot length in relation to: 1. Number of resources (buds) per shoot; 2. Success in establishment of larvae in galls; 3. Gall size and resources per gall; and 4. Survival of larvae after establishment as influenced by plant resistance and natural enemy attack. The most important factors proved to be success in establishment of larvae, with percent of variance accounted for ranging from 57 to 77 percent in three of four sites where relationships were significant, and survival after establishment of larvae, with variance accounted for ranging from 40 to 54 percent in the same three sites. The pattern of survival was dictated by plant resistance and not by natural enemies. These two additive factors resulted in a general relationship across all sites of increasing emergence of fully developed larvae per cohort as shoot length increased, accounting for 78 percent of the variance. These adaptive advantages to attacking longer shoots are sufficient to account for the pattern of increased probability of shoots being attacked as they increase in length.     

Regional tectonics, disturbance, and population size of isolated stands of Nothofagus fusca (Nothofagaceae) in a forest ecotone in south-western New Zealand

Aim New Zealand's cool temperate forests are usually dominated by one or more of the five native taxa of Nothofagus (Nothofagaceae; southern beech), but in certain regions there are sharp boundaries against podocarp–broadleaved forest where Nothofagus is rare or completely absent, either for historical (Pleistocene Glaciation) or climatic/biological (mild superhumid climate and competition) reasons. The dynamics of a Nothofagus boundary was investigated by monitoring disturbance-initiated establishment of isolated stands of N. fusca at the extreme limits of its regional distribution. Location The research was carried out in a regional forest ecotone between Nothofagus forest and podocarp–broadleaved forest in the upper Taramakau Valley, South Island, New Zealand. The survey region straddles a major, active fault system and associated tectonic movements and earthquakes with more distant epicentres have contributed to intermittent canopy disturbance of the local forests. Methods Isolated stands of Nothofagus fusca beyond the limits of continuous Nothofagus forest were investigated during two field surveys, separated by 7–10 years. Changes in population size, stem diameter of individual trees, stand basal area and mean annual diameter increment were calculated for each of fifty-four isolated stands. Types of past and recent disturbance and the probable cause of mortality of trees were noted. Results The total population of fifty-four sample stands, ranging in size from one to > 400 stems, increased by 37.4%, and compound basal area increased by 4.7% between the two surveys. Mean stem diameter growth of isolated stands was lower than expected by empirical data for N. fusca, suggesting reduced wood increment at the limits of its distribution. Tree mortality was 0.8% per year. Fifty-one per cent of the dead stems had died as a consequence of various types of natural disturbance, uproots being more common than snaps and crown breakage. Main conclusions The isolated N. fusca stands preferentially occupy sites likely to experience intermittent disturbance, mostly including disturbance of the soil cover, which facilitates their initial establishment and persistence. Because of causal relationships between mass movement on steep slopes and erosion/deposition of talus fans and river terraces, disturbance-initiated changes in forest composition are observed across a range of different landforms.     

Live crown tops of Japanese cedar (Cryptomeria japonica) approach one another in height at urban forest sites

Since the 1980s, we have found Japanese cedar, Cryptomeria japonica (L. f.) D. Don, trees with crown tops affected by dieback in isolated urban forests on the low altitude plain. To clarify future growth of C. japonica in these forests, we investigated their growth, decline levels and water status. The live crown-top heights from the ground (LCTHs) per diameter at breast height (dbh) were lower in forests with C. japonica top dieback than in forests with no top dieback. Furthermore, in a forest with C. japonica top dieback, the LCTHs were similar between trees although dbh and decline levels were different. Moreover, water status near the top of the live foliage was very similar although decline levels were different, suggesting that in urban forests, where C. japonica top dieback is observed, the LCTH is subject of restriction. A sudden increase in temperatures since the 1980s may be restricting the LCTHs of C. japonica in urban forests. We conclude that LCTHs of C. japonica in urban forests are becoming lower and more equal in each forest. If temperature continues to increase, height restriction will become more severe and LCTHs of C. japonica in urban forests will become even lower.     

Composition and ecology of vascular epiphyte communities along an altitudinal gradient in central Veracruz,Mexico

Abstract. Vascular epiphytes were studied in forests at altitudes from 720 to 2370 m on the Atlantic slope of central Veracruz, Mexico. The biomass of all trees of each species > 10 cm diameter at breast height within plots between 625 and 1500 m² was estimated. The number of species per plot ranged between 22 and 53, and biomass between 9 and 249 g dry weight/m². The highest values, both of species and biomass, were found at an intermediate altitude (1430 m). Habitat diversity may contribute to epiphyte diversity in humid forests, but the importance of this effect could not be distinguished from the influence of climate. A remarkably high number of bromeliads and orchids grew in relatively dry forests at low altitudes. In wet upper montane forests, bromeliads were replaced by ferns, while orchids were numerous at all sites, except for a pine forest. The number of epiphytic species and their biomass on a tree of a given site were closely related to tree size. According to Canonical Correspondence Analysis, the factor determining the composition of the epiphytic vegetation of a tree was altitude and to some extent tree size, whereas tree species had practically no influence. The only trees which had an evidently negative effect on epiphytes were pines, which were particularly hostile to orchids and to a lesser degree to ferns, and Bursera simaruba, which generally had few epiphytes due to its smooth and defoliating bark.     

Ecological wood anatomy of NepaleseRhododendron (Ericaceae). 1. Interspecific variation

The relationship of selected wood anatomical characters of NepaleseRhododendron with stem diameter, plant height, altitude, and plant form was investigated. We studied one to three specimens each of 26 species: five species each of trees and subtrees I, three species of subtrees II, and 13 species of shrubs. Multiple regression analysis and actual distribution of character values show that pore characters and multiseriate ray ratio have a stronger correlation with stem diameter than altitude; that pore density, vessel element length, fiber-tracheid length, and multiseriate ray density and width are equally related to altitude and stem diameter, or to altitude and plant height; and that bar number, and multiseriate ray area and height have a stronger connection with altitude. Among the characters, average pore area is most strongly correlated with stem diameter and increases exponentially as diameter increases. For wood structure of NepaleseRhododendron, 17 to 63 % of the variation is affected by non-anatomical factors. The general trends in wood structure of NepaleseRhododendron show that trees and subtrees form one continuous unit whereas shrubs form another that often has wider ranges of variation.     

Patterns and monitoring of Sesia apiformis infestations in poplar plantations at different spatial scales

Poplar plantations are expanding worldwide but little is known about the hornet clearwing moth, Sesia apiformis, one of their most severe pests. Thirty‐two poplar plantations of I‐214 clone were sampled in Spain, according to a factorial design combining stand age, site condition and understorey management to investigate the main drivers of S. apiformis habitat selection at both tree and stand level. In each plantation, one pheromone trap was activated during the flight season to test the correlation between captures and percentage of attacked trees. The proportion of other poplar forests in surrounding landscapes was calculated and used as a covariate in predictive models of trap catches. There were significantly more attacked trees in older stands. In young stands, the percentage of infested trees increased with the percentage cover of understorey vegetation. There was no significant effect of site quality on the rate of infestation and no difference in tree height, diameter or crown condition between attacked and un‐attacked trees within each stand, suggesting that S. apiformis could behave as primary pest. We hypothesized that the critical stage in the life cycle of the moth was the first instar larvae, which may benefit from protection of deep bark cracks in older stands and understorey vegetation in younger stands. We observed a positive correlation between trap capture and percentage of attacked trees in a radius of 100 m around the trap. The regression of trap catches against percentage of attacked trees was improved when the area of other poplar plantations within a distance of 600 m was incorporated in the model. This suggests that surrounding poplar stands may act as sources of immigrating moths in monitored stands. Our findings confirm that S. apiformis should be considered as a potential threat to poplar plantations and that pheromone trapping provides a suitable monitoring tool.     

Effects of an Epiphytic Orchid on Arboreal Ant Community Structure in Panama

Epiphytes are conspicuous structural elements of tropical forest canopies. Individual tree crowns in lowland forests may support more than 30 ant species, yet we know little about the effects of epiphytes on ant diversity. We examined the composition of arboreal ant communities on Annona glabra trees and their interactions with the epiphytic orchid Caularthron bilamellatum in Panama. We surveyed the ants on 73 trees (45 with C. bilamellatum and 28 lacking epiphytes) and recorded their nest sites and behavioral dominance at baits. We found a total of 49 ant species (in 20 genera), ranging 1–9 species per tree. Trees with C. bilamellatum had higher average (±SD) ant species richness (4.2±2.28) than trees without epiphytes (2.7±1.21). Hollow pseudobulbs (PBs) of C. bilamellatum were used as nest sites by 32 ant species, but only 43 percent of suitable PBs were occupied. Ant species richness increased with PB abundance in trees, but nest sites did not appear to be a limiting resource on A. glabra. We detected no close association between ants and the orchid. We conclude that higher ant species richness in the presence of the orchid is due to bottom‐up effects, especially the year‐round supply of extrafloral nectar. The structure of ant communities on A. glabra partly reflects interference competition among behaviorally dominant species and stochastic factors, as observed in other forests.     

Thinning Jeffrey pine stands to reduce susceptibility to bark beetle infestations in California,U.S.A.

• 1 Bark beetles (Coleoptera: Curculionidae, Scolytinae) are commonly recognized as important tree mortality agents in coniferous forests of the western U.S.A.
• 2 High stand density is consistently associated with bark beetle infestations in western coniferous forests, and therefore thinning has long been advocated as a preventive measure to alleviate or reduce the amount of bark beetle‐caused tree mortality.
• 3 The present study aimed to determine the effectiveness of thinning to reduce stand susceptibility to bark beetle infestations over a 10‐year period in Pinus jeffreyi forests on the Tahoe National Forest, California, U.S.A. Four treatments were replicated three times within 1‐ha square experimental plots. Treatments included thinning from below (i.e. initiating in the smallest diameter classes) to a residual target basal area (cross‐sectional area of trees at 1.37 m in height) of: (i) 18.4 m²/ha (low density thin); (ii) 27.6 m²/ha (medium density thin); (iii) 41.3 m²/ha (high density thin); and (iv) no stand manipulation (untreated control).
• 4 Throughout the present study, 107 trees died as a result of bark beetle attacks. Of these, 71% (75 trees) were Abies concolor killed by Scolytus ventralis; 20.6% (22 trees) were Pinus ponderosa killed by Dendroctonus ponderosae; 4.7% (five trees) were P. jeffreyi killed by Dendroctonus jeffreyi; 1.8% (two trees) were P. jeffreyi killed by Ips pini; 0.9% (one tree) were P. jeffreyi killed by Orthotomicus (= Ips) latidens; 0.9% (one tree) were P. ponderosa killed by both Dendroctonus brevicomis and D. ponderosae; and 0.9% (one tree) were P. jeffreyi killed by unknown causes.
• 5 In the low density thin, no pines were killed by bark beetles during the 10‐year period. Significantly fewer trees (per ha/year) were killed in the low density thin than the high density thin or untreated control. No significant treatment effect was observed for the percentage of trees (per year) killed by bark beetles.

     

Stem-boring insects of fragmented Calamagrostis habitats: herbivore–parasitoid community structure and the unpredictability of grass shoot abundance

Abstract.1. Communities of stem-boring insects attacking the grass Calamagrostis epigeios (L.) Roth were analysed from twenty-five pure stands of C. epigeios differing in area and isolation. Insect communities comprised nine phytophagous and eighteen entomophagous species. Most abundant herbivores (attacking > 1% of all shoots) were Tetramesa eximia (Giraud) (Hym. Eurytomidae), Eurytoma sp. (Hym. Eurytomidae), Lasioptera calamagrostidis (Rübs.) (Dipt. Cecidomyiidae), and Eriopeltis sp. (Hom. Pseudococcidae). About 28% of the twenty-seven species were monophagous, 61% oligophagous (restricted to Poaceae), and 11% polyphagous. Although herbivores generally attacked thick shoots, species-specific differences in the attacked mean diameter were found.2. The main purpose of this study was to analyse the relative effects of area, isolation and unpredictability of available food resources on the abundance of herbivores, parasitoids and resulting tritrophic effects. Variability of shoot abundance varied greatly between years, and even more between habitats within the same year. On average, shoot density per habitat increased or decreased 2.4-fold between succeeding years.3. Variability of grass shoot abundance was not related to habitat characteristics such as area, isolation, biomass, shoot density, changes in area or shoot density between years, or shoot features like internode number or diameter, but was negatively influenced by shading of trees. Similarly, habitat area or isolation did not correlate with other habitat features.4. Grass shoot abundance was the most important predictor of species abundance and attack rate of herbivores. In contrast to expectations, area and isolation of habitats did not explain differences in species richness, and only influenced abundance of three species. In addition, even the experimental removal of all grass shoots (and thereby, of all stem-boring insects) from five habitats in winter did not negatively influence colonization the following year. These results suggest a surprisingly high mobility of this rather specialized community of stem-boring insects that appeared to superimpose on area and isolation effects. Unpredictability of grass shoot abundance was presumably the main selection pressure for the obviously high dispersal abilities of the Calamagrostis insects.5. Per cent parasitism was mainly influenced by the percentage of attacked shoots. Parasitism of T. eximia by its major parasitoid also correlated with habitat connectivity, emphasizing the hypothesis that a continuum of habitats should be particularly favourable for the third trophic level.