Dynamics of macronutrients during the first stages of litter decomposition from forest species in a temperate area (Galicia,NW Spain)

The decomposition and dynamics of nutrient elements was studied for leaves and twigs from Quercus robur, Pinus radiata and Eucalyptus nitens in soils developed from granodiorite, slate and limestone in Galicia, NW Spain. Two 1-ha plots were selected for each material-vegetation combination, making a total of 18 plots. Litter decomposition and nutrient dynamics during the first six months were investigated using litterbags. Mass loss was higher for leaves than twigs and not significantly different for different tree species. Mass loss correlated significantly with carbon loss (r = 0.96 for leaves, r = 0.90 for twigs). As a general trend, nutrient release from leaves and twigs was greater for the broadleaved species (eucalyptus and oak). K and S were rapidly released from all litters, while Ca and Mg showed the highest tendency to be immobilised. N and P behave similarly, with a final balance of net release. Soil parent material did not significantly influence decomposition or nutrient dynamics.     

Changes in Heartwood Chemistry of Dead Yellow-Cedar Trees that Remain Standing for 80 Years or More in Southeast Alaska

We measured the concentrations of extractable bioactive compounds in heartwood of live yellow-cedar (Chamaecyparis nootkatensis) trees and five classes of standing snags (1–5, averaging 4, 14, 26, 51, and 81 years-since-death, respectively) to determine how the concentrations changed in the slowly deteriorating snags. Three individuals from each of these six condition classes were sampled at four sites spanning a 260-km distance across southeast Alaska, and the influence of geographic location on heartwood chemistry was evaluated. Cores of heartwood were collected at breast height and cut into consecutive 5-cm segments starting at the pith. Each segment was extracted with ethyl acetate and analyzed by gas chromatography. Concentrations of carvacrol, nootkatene, nootkatol, nootkatone, nootkatin, and total extractives (a sum of 16 compounds) for the inner (0–5 cm from pith), middle (5–10 cm from pith), and surface (outer 1.1–6.0 cm of heartwood) segments from each core were compared within each tree condition class and within segments across condition classes. Heartwood of class 1 and 2 snags had the same chemical composition as live trees. The first concentration changes begin to appear in class 3 snags, which coincides with greater heartwood exposure to the external environment as decaying sapwood sloughs away, after losing the protective outer bark. Within core segments, the concentrations of all compounds, except nootkatene, decrease between snag classes 2 and 5, resulting in the heartwood of class 5 snags having the lowest quantities of bioactive compounds, although not different from the amounts in class 4 snags. This decline in chemical defense is consistent with heartwood of class 5 snags being less decay-resistant than heartwood of live trees, as observed by others. The unique heartwood chemistry of yellow cedar and the slow way it is altered after death allow dead trees to remain standing for up to a century with a profound impact on the ecology of forests in southeast Alaska where these trees are in decline.     

Volatiles from Potential Hosts of Rhopalicus tutela a Bark Beetle Parasitoid

Host location cues for parasitic wasps that attack bark beetle larvae concealed under the bark of spruce trees were analyzed by collecting odor samples from entrance holes into the bark beetle galleries, isolated larvae, and pupal chambers with or without bark beetle larvae. Odor samples were collected by dynamic headspace adsorptions on Porapak Q or static adsorptions by using solid-phase microextraction (SPME) with Carbowax–divinylbenzene as the adsorbing phase. Samples were analyzed by coupled gas chromatographic–electroantennographic detection (GC-EAD) and GC-mass spectrometry (GC-MS). The antennae of Rhopalicus tutela females responded primarily to oxygenated monoterpenes that are typical for damaged host trees. These compounds are attractive to bark beetle parasitoids in long-range host location, suggesting that they are used in both long- and short-range host location. No differences could be detected between samples collected from pupal chambers with or without mature larvae. Larvae outside pupal chambers emitted low quantities of the same compounds present in empty pupal chambers. The data support the hypothesis that volatiles used by host foraging parasitoids arise from the interaction between introduced microorganisms and the bark and/or vascular tissue of the host tree rather than from the bark beetle larvae.     

Primary Attraction of the Fir Engraver, Scolytus ventralis

In laboratory bioassays, Porapak Q-captured and steam-distilled volatiles from the bark of host trees, Abies grandis, particularly from root-rot-infected trees, attracted 50–70% of male and female fir engravers, Scolytus ventralis. Gas chromatographic–electroantennographic detection (GC-EAD) analyses of Porapak Q-captured bark volatiles revealed 19 EAD-active compounds of which 13 (mostly monoterpenes) were identified by GC–mass spectrometry (GC-MS). In separate field experiments, multiple-funnel traps baited with two blends of these 13 synthetic volatiles released at 280 and 340 mg/ 24 hr attracted 66 and 93% of the total S. ventralis captured, respectively. The clerid predator, Thanasimus undulatus, also responded strongly to the kairomonal volatiles. Additional experiments produced no evidence for aggregation pheromones in S. ventralis. These included laboratory bioassays and GC and GC-EAD analyses of Porapak Q-captured volatiles from male- and female-infested logs or trees undergoing mass attack in the field, GC analyses and/or bioassays of extracts from female accessory glands, extracted volatiles from emerged, attacking and juvenile hormone-treated beetles of both sexes, and videotape analysis of the behavior of attacking beetles on the bark surface. We argue against the hypothesis of pheromone-mediated secondary attraction in S. ventralis and conclude that the attack dynamics of this species can be explained solely by its sensitive primary attraction response to host volatiles.     

Defensive mechanisms of loblolly and shortleaf pine against attack by southern pine beetle,Dendroctonus frontalis Zimmermann,and its fungal associate,Ceratocystis minor (Hedgecock) Hunt

Loblolly and shortleaf pine growing on a single site in the North Carolina piedmont were examined to determine similarities and differences in their defensive mechanisms against the southern pine beetle,Dendroctonus frontalis Zimmermann, and its fungal associate,Ceratocystis minor (Hedgecock) Hunt. Both species responded to wounding and fungal inoculation by forming a hypersensitive lesion around the wound site. There were significantly less soluble sugars and more monoterpenes in the lesion tissue than in unwounded inner bark. The two species were similar in resin flow rate and inner bark soluble sugar content, but the loblolly pines had thicker bark, longer hypersensitive lesions, and a higher concentration of inner bark monoterpenes. Inner bark monoterpene composition was also significantly different between the two pine species. It is hypothesized that two different defensive strategies against southern pine beetle attack may be utilized.     

Host selection behavior of bark beetles (Coleoptera: Scolytidae) attackingPinus ponderosa,with special emphasis on the western pine beetle,Dendroctonus brevicomis

Detection of weakened hosts from a distance by bark beetles through olfaction was investigated in field experiments. No significant numbers of Scolytidae were attracted to anaerobically treated pine bolts, stem disks, or sugar and ponderosa pine bark including phloem. Treatment of living trees with cacodylic acid induced attacks byDendroctonus brevicomis, D. ponderosae, Ips latidens, Gnathotrichus retusus, andPityophthorus scalptor, beginning two weeks after treatment. There was no significant difference between landing rates ofD. brevicomis andD. ponderosae on screened treated trees and screened controls. There was a significant increase in landing rates ofG. retusus andI. latidens, because both species had penetrated the screen and produced pheromones. Tree frilling alone did not increase the landing rate of bark beetles. Freezing of the lower trunk with dry ice did not increase significantly the landing rate ofD. brevicomis, D. ponderosae, G. retusus, orI. latidens on screened trees, whereas unscreened frozen trees were attacked by all four species. There was no significantly higher landing rate byD. brevicomis, D. ponderosae, I. paraconfusus, I. latidens, G. retusus, orHylurgops subcostulatus on screened trees evidencing symptoms of severe infection by the root pathogenVerticicladiella wagenerii, than on symptornless trees. These experiments show thatD. brevicomis, D. ponderosae, I. paraconfusus, I. latidens, andG. retusus land, apparently indiscriminately, on healthy and stressed hosts. Thus, in these species host discrimination must occur after landing and prior to sustained feeding.These studies were supported in part by the U.S.D.A. Forest Service, Cooperative State Research Services (2598-RRF.W-1 10) and the National Science Foundation and Environmental Protection Agency through a grant (NSF GB-34719/BM575-04223) to the University of California; and by the Canada Department of the Environment. The findings, opinions and recommendations are not necessarily those of the University of California or the funding agencies.From a thesis submitted by H.A. Moeck to the University of California, Berkeley, in partial fulfillment of the requirements for the degree Doctor of Philosophy in Entomology.     

Influence of aggregation inhibitors (verbenone and ipsidenol) on landing and attack behavior ofDendroctonus brevicomis (Coleoptera: Scolytidae)

The influence of the aggregation inhibitors verbenone and ipsdienol on the response of western pine beetle,Dendroctonus brevicomis, to attractive host trees was investigated. Paired ponderosa pine trees (Pinus ponderosa) were baited with aggregation semiochemicals to stimulate mass attack. One tree in each pair received an inhibitor treatment consisting of five sets of two verbenone and two ipsdienol dispensers spaced 1 m apart vertically along the tree bole. Beetle landing was monitored with sticky traps on the tree bole, and attack density was assessed from bark samples removed four or seven days after baiting. The inhibitor treatment resulted in a significant reduction of both the numbers of beetles landing on trees and the density of attacking beetles compared to control trees (without inhibitors). The ratios of beetle landing density to attacking density were not different between inhibitor-treated and control trees, nor were the vertical distributions of beetles landing or attacking, suggesting that beetle behavior was primarily influenced at a longer range, prior to landing on the tree. Although the application of verbenone and ipsdienol did not preventD. brevicomis from attacking baited trees, our results suggest that when applied to unattacked (and unbaited) trees, their effectiveness at reducing the attack pressure might allow trees having a certain amount of resistance to survive attack by pioneer beetles.     

Behavioral responses of elm bark beetles to baited and unbaited elms killed by cacodylic acid

Diseased elms, treated with various doses of cacodylic acid in northwest England, became attractive to elm bark beetles (Coleoptera: Scolytidae). This attraction seemed to be independent of pheromone baits. However attractive the trees became, they were unsuitable to the beetles as breeding sites since significantly more beetles visited the trees than were stimulated to penetrate and attempt to breed. It seems as if colonization of trap trees by the bark saprophytePhomopsis oblonga following cacodylic acid treatment made the trees unsuitable to beetles for breeding.     

Phloeophagous and predaceous insects responding to synthetic pheromones of bark beetles inhabiting white spruce stands in the Great Lakes region

Tree killing and saprophytic bark beetles exert important ecological and economic roles in North American spruce forests. Chemical signaling among bark beetles, and responses by associate insects such as predators and competitors, have significant effects on the population dynamics and ecology of this community. Synthetic pheromones of primary (tree killing) and secondary (saprophytic) bark beetle species and blank controls were tested using multiple funnel and lower stem flight traps in white spruce forests in Wisconsin, Michigan, and Minnesota. Six phloeophagous and four predaceous species were collected with significant attraction by the bark beetles Dryocoetes affaber, Dryocoetes autographus, and Polygraphus rufipennis, and the predatory checkered beetles (Coleoptera: Cleridae) Thanasimus dubius and Enoclerus nigrifrons. In general, trap catches to synthetic lures resembled the species composition obtained by felling trees and collecting emerging beetles in a companion study, although several species showed differing trends. Some cross attraction occurred among bark beetles and between bark beetles and predatory beetles. For example, P. rufipennis was abundant in traps baited with Dryocoetes spp. pheromones. Thanasimus dubius and E. nigrifrons were collected in significant numbers in traps baited with the pheromone of the spruce beetle (Dendroctonus rufipennis), frontalin plus -pinene. This is a new observation for E. nigrifrons. Attraction of T. dubius to the pheromones of at least three bark beetle species in the Great Lakes region, as well as to several southern and western species, reflects its role as a habitat specialist and feeding generalist. Several other important predators and competitors commonly obtained in pine forests in this region were not obtained in these spruce stands, either in response to synthetic pheromones of spruce colonizing beetles, or in host material colonized by these beetles. Potential differences in predator prey dynamics between spruce and pine ecosystems in the Great Lakes region are discussed.     

Trap trees for elm bark beetles

European elm bark beetles,Scolytus multistriatus (Marsh.), were strongly attracted to American elms,Ulmus americana L., baited with theS. multistriatus attractant, multilure, or killed by injection of the arboricide, cacodylic acid; a combination of the two treatments was most attractive. Comparisons of beetle catches on sticky bands affixed to the trees with samples of bark from the same trees showed that the number of beetles landing on cacodylic acid-treated trees was approximately 40 times greater than the number boring into them. Spraying the bark with the insecticide chlorpyrifos had no direct effect on attraction. No live bark beetle brood was found in trees that had been treated with cacodylic acid or chlorpyrifos, but trees that were only baited or left untreated (check) were attacked, killed, and colonized. We suggest that the contribution of the cacodylic acid trap tree technique to Dutch elm disease control will be enhanced by baiting treated trees with multilure and spraying their lower boles with 0.5% chlorpyrifos. This treatment will eliminate diseased and unwanted elms as potential breeding material and kill large numbers of elm bark beetles that might otherwise innoculate healthy elms with the Dutch elm disease fungus.     

Relationships betweenDendroctonus micans Kug. (Coleoptera: Scolytidae) survival and development and biochemical changes in Norway Spruce,Picea abies (L.) Karst., phloem caused by mechanical wounding

The bark of Norway spruce trees,Picea abies (L.) Karst., was wounded to produce areas of bark with differing biochemical characteristics. Adults and larvae ofDendroctonus micans Kug. were inserted into these trees at points around the wounds. Larval survival was higher, larval dry weights were higher, and adults were more likely to oviposit at sites around the wound that had higher phloem moisture contents. Larvae showed higher survival and dry weights in phloem with low terpene contents. Adults oviposited in phloem with lower stilbene glucoside contents. Phloem moisture differed at sites around the wound compared with that at a control site away from the wound, and the nitrogen content of entire bark cores was higher above wounds than at control sites away from the wounds. Phloem moisture correlated with the nitrogen content of entire bark. Phloem terpene contents correlated with each other, as did the stilbene contents. Associations between the host exploitation pattern ofD. micans and the biochemistry of the host phloem are discussed in relation to preselection by adults of sites suitable for larval development.     

Antifeedants against <Emphasis Type="Italic">Hylobius abietis</Emphasis> Pine Weevils: An Active Compound In Extract of Bark of <Emphasis Type="Italic">Tilia cordata</Emphasis> Linden

Linden (Tilia cordata) bark was shown to contain an antifeedant effective against the large pine weevil, Hylobius abietis. Soxhlet extraction of inner and outer bark resulted in an extract that showed antifeedant activity in a microfeeding assay. The extract was fractionated by chromatography on silica gel using gradient elution with solvents of increasing polarity. The content of the fractions obtained was monitored by thin layer- and gas chromatography. Fractions of similar chemical composition were merged. Two of the 17 fractions showed antifeedant activity in the microfeeding assay. Nonanoic acid was identified in both of these fractions. Subsequent testing in the microfeeding assay showed that nonanoic acid possessed strong antifeedant activity against H. abietis adults.     

Resistance of western white pine to feeding and oviposition byPissodes strobi peck in western canada

Forced and choice feeding and oviposition experiments were conducted by caging overwinteredPissodes strobi on lateral branches or leaders of sympatric Engelmann spruce and/or western white pine trees in Manning Provincial Park, British Columbia, or on transplanted, potted trees in Burnaby, British Columbia. Whereas femaleP. strobi preferred to feed on 1-year-old Engelmann spruce lateral branches in both the forced- and choice-feeding experiments, male weevils failed to discriminate between this host and western white pine. Forced-feeding experiments that utilized 1-year-old leaders demonstrated that western white pine is an acceptable host forP. strobi feeding, but that the native host, Engelmann spruce, is preferred under choice conditions. In all experiments, female weevils oviposited only on Engelmann spruce. These results indicate that the releasing stimulus for feeding is present in the bark of both conifer species and suggests that a separate releasing stimulus for oviposition is absent from western white pine.Coleoptera: Curculionidae.Research supported by Canadian Forestry Service Science Subvention Grant, National Science Foundation (USA) Grant No. BMS-7413643, and National Research Council (Canada) Grant No. A3881.     

Semiochemicals from bark beetles: New results,remarks, and reflections

A brief survey is given about recent results in the identification of semiochemicals in bark beetles: Males ofIps sexdentatus (Boern.), stressed by the attack on resinous trees produce large amounts of 3(S)-1-methyl-5-(1-hydroxyl-1-methylethyl)-cyclohexa-1,3-diene. The compound appears to be derived from ³-carene and acts as a repellent. Males ofIps typographus (L.), stressed through the attack on unsuitable host material release 3-methyl-7-methylene-1,3(E), 8-nonatriene, which seems to act as a repellent. The odor bouquet of three species ofPityogenes is described. The occurrence of (+)-grandisol and other compounds related to weevil pheromones points to a close relation between Scolytidae and Curculionidae. Females ofDendroctonus simplex (Le Conte) use (–)-frontalin as the main pheromone. 6-Methyl-6-hepten-2-one, a minor component among the volatile compounds released by the females, is regarded as a possible precursor of frontalin. Similarly, (2R,5S)-2(1-hydroxyl-1-methylethyl)-5-methyltetrahydrofuran, pityol, a pheromone ofPityophthorus spp., is regarded to at least share a common biogenetic precursor with 6-methyl-5-hepten-2-ol, sulcatol. A new bicylic acetal, 2-ethyl-1,5-dimethyl-6,8-dioxabicyclo[3.2.1]octane, is described as an aggregation pheromone of the beech bark beetle,Taphrorychus bicolor (Herbst). Structural relationships between bark beetle pheromones and plant volatiles are discussed.     

Phenolics in black oak bark and leaves

Catechin, quercitrin, robinin, quercetin 3-methyl ether, scopoletin, cholorogenic acid, several leucoanthocyanins, and condensed and hydrolyzable tannins were identified in bark and leaves ofQuercus velutina Lamarck. The concentrations of most phenolics in leaves increased as the growing season progressed, whereas those of most phenolics in bark remained essentially unchanged. Qualitative differences in bark and leaf phenolics among different trees were negligible.     

California Ground Squirrel (<Emphasis Type="Italic">Spermophilus beecheyi</Emphasis>) Defenses against Rattlesnake Venom Digestive and Hemostatic Toxins

Previous studies have shown that some mammals are able to neutralize venom from snake predators. California ground squirrels (Spermophilus beecheyi) show variation among populations in their ability to bind venom and minimize damage from northern Pacific rattlesnakes (Crotalus oreganus), but the venom toxins targeted by resistance have not been investigated. Four California ground squirrel populations, selected for differences in local density or type of rattlesnake predators, were assayed for their ability to neutralize digestive and hemostatic effects of venom from three rattlesnake species. In Douglas ground squirrels (S. b. douglasii), we found that animals from a location where snakes are common showed greater inhibition of venom metalloprotease and hemolytic activity than animals from a location where snakes are rare. Effects on general proteolysis were not different. Douglas ground squirrels also reduced the metalloprotease activity of venom from sympatric northern Pacific rattlesnakes (Crotalus oreganus oreganus) more than the activity of venom from allopatric western diamondback rattlesnakes (C. atrox), but enhanced fibrinolysis of sympatric venom almost 1.8 times above baseline levels. Two Beechey ground squirrel (S. b. beecheyi) populations had similar inhibition of venoms from northern and southern Pacific rattlesnakes (C. o. helleri), despite differences between the populations in the locally prevalent predator. However, the venom toxins inhibited by Beechey squirrels did vary among venom from Pacific rattlesnake subspecies, and between these venoms and venom from allopatric western diamondback rattlesnakes. Blood plasma from Beechey squirrels showed highest inhibition of metalloprotease activity of northern Pacific rattlesnake venom, general proteolytic activity and hemolysis of southern Pacific rattlesnake venom, and hemolysis by allopatric western diamondback venom. These results reveal previously cryptic variation in venom activity against resistant prey that suggests reciprocal adaptation at the molecular level.     

Oviposition stimulants for the beetle,Monochamus alternatus hope,in inner bark of pine

Field and laboratory ovipositional responses ofMonochamus alternatus Hope, respectively, to methanol and water extracts from pine inner bark were examined in comparison with those to pine inner bark, especially using a laboratory-built apparatus for the latter bioassay. Irrespective of the existence of volatiles from paraquat-induced lightwood, pine inner bark and its methanol and water extracts stimulated ovipositional response only in the presence of free moisture.     

Aqueous Leachate of Lantana camara Kills Water Hyacinth

Lantana camara, a terrestrial plant, was screened for allelopathic impact on water hyacinth (Eichhornia crassipes). Water hyacinth was allowed to grow in experimental pots containing 3% aqueous leachate (w/v) of Lantana twigs. The leachate was allelopathic to the growth of water hyacinth, and killed water hyacinth after 21 days under the experimental conditions. Leachate concentrations from 1–3% of Lantana were highly toxic to water hyacinth plants. Leachate from young Lantana twigs with prickly orange, pink, and yellow flowers (multicolored) was more toxic than leachate from mature twigs. This study indicates the potential for utilization of terrestrial allelopathic species such as Lantana camara to suppress the growth of water hyacinth.     

Bark consumption by voles in relation to mineral contents

Recent field experiments with impregnated wooden sticks have demonstrated a pronounced use by small rodents of mineral supplies, especially sodium, and such findings seemed related to vole damage to forestry seedlings. Consumption of the bark of experimentally introduced aspen twigs and of sodium-impregnated sticks by voles (mainly or onlyMicrotus agrestis) correlated significantly on clear-cuts but not on unmanipulated abandoned fields. Such a correlation appeared when abandoned fields were cut continuously in summer. At vole peak densities, bark of pine seedlings experimentally fertilized with sodium was consumed but not bark of seedlings fertilized with calcium or control seedlings. Field pine seedlings attacked by voles had significantly higher levels of calcium, sodium, and phosphorus than the nearest untouched seedling. However, sodium and phosphorus contents correlated strongly. Sodium and calcium supply to voles in laboratory feeding trials did not diminish the moderate interest in pine bark. Such conditions are, however, assumed to mimic a situation of bark sampling in low-density populations. Sodium, and possibly also calcium, requirements are concluded to be partial determinants of the destructive bark consumption by voles at the peaks of their multiannual population cycles.     

California Ground Squirrel (Spermophilus beecheyi) Defenses Against Rattlesnake Venom Digestive and Hemostatic Toxins

Previous studies have shown that some mammals are able to neutralize venom from snake predators. California ground squirrels (Spermophilus beecheyi) show variation among populations in their ability to bind venom and minimize damage from northern Pacific rattlesnakes (Crotalus oreganus), but the venom toxins targeted by resistance have not been investigated. Four California ground squirrel populations, selected for differences in local density or type of rattlesnake predators, were assayed for their ability to neutralize digestive and hemostatic effects of venom from three rattlesnake species. In Douglas ground squirrels (S. b. douglasii), we found that animals from a location where snakes are common showed greater inhibition of venom metalloprotease and hemolytic activity than animals from a location where snakes are rare. Effects on general proteolysis were not different. Douglas ground squirrels also reduced the metalloprotease activity of venom from sympatric northern Pacific rattlesnakes (C. o. oreganus) more than the activity of venom from allopatric western diamondback rattlesnakes (C. atrox), but enhanced the fibrinolysis of sympatric venom almost 1.8 times above baseline levels. Two Beechey ground squirrel (S. b. beecheyi) populations had similar inhibition of venoms from northern and southern Pacific rattlesnakes (C. o. helleri), despite differences between the populations in the locally prevalent predator. However, the venom toxins inhibited by Beechey squirrels varied among venom from Pacific rattlesnake subspecies, and between these venoms and venom from allopatric western diamondback rattlesnakes. Blood plasma from Beechey squirrels showed highest inhibition of metalloprotease activity of northern Pacific rattlesnake venom, general proteolytic activity and hemolysis of southern Pacific rattlesnake venom, and hemolysis by allopatric western diamondback venom. These results reveal previously cryptic variation in venom activity against resistant prey that suggests reciprocal adaptation at the molecular level.