Biologically induced mineralization in the tree Milicia excelsa (Moraceae): its causes and consequences to the environment

Iroko trees (Milicia excelsa) in Ivory Coast and Cameroon are unusual because of their highly biomineralized tissues, which can virtually transform the trunk into stone. Oxalic acid (C₂O₄H₂) and metal‐oxalate play important roles in their ecosystems. In this study, the various forms of oxalate and carbonate mineralization reactions are investigated by using scanning electron microscopy and X‐ray diffraction. Calcium oxalate monohydrate is associated with stem, bark and root tissues, whereas calcium oxalate dihydrate is found with wood rot fungi in soils, as well as in decaying wood. Laboratory cultures show that many soil bacteria are able to oxidize calcium oxalate rapidly, resulting in an increase in solution pH. In terms of M. excelsa, these transformations lead to the precipitation of calcium carbonate, not only within the wood tissue, but also within the litter and soil. We calculate that c. 500 kg of inorganic carbon is accumulated inside an 80‐year‐old tree, and c. 1000 kg is associated with its surrounding soil. Crucially, the fixation of atmospheric CO₂ during tree photosynthesis, and its ultimate transformation into calcite, potentially represents a long‐term carbon sink, because inorganic carbon has a longer residence time than organic carbon. Considering that calcium oxalate biosynthesis is widespread in the plant and fungal kingdoms, the biomineralization displayed by M. excelsa may be an extremely common phenomena.     

Fungi,bacteria and soil pH: the oxalate–carbonate pathway as a model for metabolic interaction

The oxalate–carbonate pathway involves the oxidation of calcium oxalate to low‐magnesium calcite and represents a potential long‐term terrestrial sink for atmospheric CO ₂. In this pathway, bacterial oxalate degradation is associated with a strong local alkalinization and subsequent carbonate precipitation. In order to test whether this process occurs in soil, the role of bacteria, fungi and calcium oxalate amendments was studied using microcosms. In a model system with sterile soil amended with laboratory cultures of oxalotrophic bacteria and fungi, the addition of calcium oxalate induced a distinct pH shift and led to the final precipitation of calcite. However, the simultaneous presence of bacteria and fungi was essential to drive this pH shift. Growth of both oxalotrophic bacteria and fungi was confirmed by qPCR on the frc (oxalotrophic bacteria) and 16S rRNA genes, and the quantification of ergosterol (active fungal biomass) respectively. The experiment was replicated in microcosms with non‐sterilized soil. In this case, the bacterial and fungal contribution to oxalate degradation was evaluated by treatments with specific biocides (cycloheximide and bronopol). Results showed that the autochthonous microflora oxidized calcium oxalate and induced a significant soil alkalinization. Moreover, data confirmed the results from the model soil showing that bacteria are essentially responsible for the pH shift, but require the presence of fungi for their oxalotrophic activity. The combined results highlight that the interaction between bacteria and fungi is essential to drive metabolic processes in complex environments such as soil.     

Oxalate accumulation from citrate by Aspergillus niger

Carbon-14 was incorporated from citrate-1,5-¹⁴C, glyoxylate-¹⁴C(U), or glyoxylate-1-¹⁴C into oxalate by cultures of Aspergillus niger pregrown on a medium with glucose as the sole source of carbon. Glyoxylate-¹⁴C(U) was superior to glyoxylate-1-¹⁴C and citrate-1,5-¹⁴C as a source of incorporation. By addition of a great amount of citrate the accumulation of oxalate was accelerated and its maximum yield increased. In a cell-free extract from mycelium forming oxalate from citrate the enzyme oxaloacetate hydrolase (EC 3.7.1.1) was identified. Its in vitro activity per flask exceeded the rate of in vivo accumulation of oxalate. Glyoxylate oxidizing enzymes (glycolate oxidase, EC 1.1.3.1; glyoxylate oxidase, EC 1.2.3.5; NAD(P)-dependent glyoxylate dehydrogenase; glyoxylate dehydrogenase, CoA-oxalylating, EC 1.2.1.17) could not be detected in cell-free extracts. It is concluded that in cultures accumulating oxalate from citrate after pregrowth on glucose, oxalate arises by hydrolytic cleavage of oxaloacetate but not by oxidation of glyoxylate.Abbreviations Used DCPIP 2,6-dichlorophenolindophenol     

A major symbiont shift supports a major niche shift in a clade of tree-killing bark beetles

1. One small clade of bark beetles, out of thousands of species worldwide, has shifted from using phloem to using a combination of phloem and outer bark, or to completely using outer bark. 2. The shift to outer bark has been accompanied by a shift to non-typical bark beetle mutualist fungi in Entomocorticium (Basidiomycota) and Ceratocystiopsis (Ascomycota). 3. This study compared the growth and metabolic capabilities of fungi associated with a nearly phloem-independent species, Dendroctonus brevicomis, with those of mutualist fungi of Dendroctonus ponderosae, a completely phloem-colonising beetle in a sister clade associated with more typical Ascomycota in Grosmannia and Ophiostoma. 4. Only the basidiomycete associated with D. brevicomis could degrade cellulose and lignin, whereas both the ascomycete and basidiomycete could grow in outer bark. Ascomycetes associated with D. ponderosae could not degrade cellulose or lignin or grow in outer bark. 5. Beetles and fungi in these mutualisms may best be considered as co-niche constructors. For niche construction, one organism must modify a resource in a way that enhances its fitness while also influencing fitness of other organisms using the resource. Here, the beetles kill the tree, transport the fungi into the tree, and modify the woody substrate for use by the fungi. They have also evolved mycangia to ensure vertical dissemination of the fungi. In turn, the fungi modify tree tissues and provision nutrients to the host and have evolved traits that support their acquisition and transport by the beetle host in mycangia.     

芋营养器官中的草酸钙晶体及其可能的生理作用

利用徒手切片,在光学显微镜下对芋(Colocasia esculenta(L.)Schott)营养器官中晶体的类型和分布进行了观察和研究,并用化学方法对晶体的化学成分进行了鉴定。结果表明,芋营养器官中的晶体为草酸钙结晶体,形态上可以分为针晶和簇晶两大类。含针晶束的异细胞有3种类型:含发射型草酸钙针晶束异细胞(存在于叶片、叶柄、块茎中),含大型草酸钙针晶束异细胞(存在于叶片、叶柄、块茎、块茎皮中),含大量草酸钙针晶的管状异细胞(仅存在于不定根中)。草酸钙针晶也有散乱分布于块茎和不定根中的。草酸钙簇晶在叶片、叶柄、块茎、块茎皮、不定根中均有分布,且叶片、叶柄、块茎皮中的簇晶比块茎和不定根中的尖锐。芋营养器官中的草酸钙晶体很可能是作为一种防御机制,防止动物的取食。     

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.

     

入侵害虫悬铃木方翅网蝽越冬后期在豫北寄主树干上的分布研究

【目的】入侵害虫悬铃木方翅网蝽近年来严重威胁我国城市行道树二球悬铃木的健康,受害严重时造成树木死亡。在越冬后期对悬铃木树干上的悬铃木方翅网蝽成虫的越冬习性开展研究,能为冬季防治积累生物学研究基础。【方法】早春随机调查河南省新乡市行道树二球悬铃木树干栓皮层外层下方隐藏的悬铃木方翅网蝽成虫,比较阳侧和阴侧虫口密度。同时记载悬铃木树干0.00~0.50 m、0.51~1.00 m、1.01~1.50 m、1.51~2.00 m 4个不同高度区段范围和树干皮层外层皲裂程度在5%以下、5%~25%、25%~50%、50%~75%、75%~95%、95%以上6个等级条件下害虫的虫口密度。【结果】栖息在悬铃木树干树皮下方的悬铃木方翅网蝽在阴面更多,阴面和阳面的虫口密度差异达极其显著水平[(F=6.63)(F0.01(1,19)=1.0132),P=0.00]。对寄主树干4个不同高度区段的虫口密度比较后,发现在树干1.0~1.5 m区段间的虫口密度最大,平均11.5头;且虫口密度在悬铃木不同高度区段间存在极显著差异[(F=26.91)(F0.01(3,56)=4.1519),P=0.00]。悬铃木树皮的不同皲裂程度对悬铃木方翅网蝽虫口密度的影响达极其显著水平[(F=31.02)(F0.01(3,56)=4.1519),P=0.00],在树皮外皮50%~75%皲裂程度条件下的虫口密度最大,平均9.8头。利用双因素方差分析方法分析树干不同高度和树皮不同皲裂等级的互作对虫口密度的分布影响达显著水平[(F=2.46)(F0.05(9,56)=2.0519),P=0.0195]。【结论】华北地区害虫秋冬防治时,在寄主悬铃木树干的最好涂白时机是在害虫全部下树后的12月上中旬进行,涂白高度以树干1.0~2.0 m范围为宜。本研究对悬铃木方翅网蝽在越冬期的预防和治理有较为重要的参考价值。     

The Endophytic Mycoflora of Bark,Leaf, and Stem Tissues of <Emphasis Type="Italic">Azadirachta indica</Emphasis> A. Juss (Neem) from Varanasi (India)

A systematic study was made of the endophytes of Azadirachta indica A. Juss (the neem tree) growing in several of its natural habitats in India. A total of 233 isolates of endophytic fungi representing 18 fungal taxa were obtained from segments of bark, stem, and leaves of this tree. Hyphomycetes (62.2%) were the most prevalent followed by the Coelomycetes (27.4%) and Mycelia Sterilia (7.7%). As mathematically determined, the maximum species richness and frequency of colonization of endophytes appeared in leaf segments rather than stem and bark tissues from each location. Endophytic colonization frequency was also greater in leaves (45.5%) than bark (31.5%). The leaf samples from all locations were nearly constant in their endophytic composition, whereas bark samples showed maximum diversity at different locations. Inter-site comparisons for endophytic diversity, however, were not significantly different with Loc1 and Loc2 having a maximum of 66.67% J _c. The smallest similarity was between Loc2 and Loc3 of 54.17% J _c. The dominant endophytic fungi isolated were Phomopsis oblonga, Cladosporium cladosporioides, Pestalotiopsis sp., Trichoderma sp, and Aspergillus sp. Genera such as Periconia, Stenella, and Drechslera are reported here for the first time as endophytes from this host plant. This report illustrates the value of sampling different tissues of a given plant in several locations to obtain the greatest species diversity of endophytes. The rich and sizeable collection of endophytic fungi from this specific plant may represent a unique source of one or more of the interesting and useful bioactive compounds normally associated with A. indica such as the azadirachtins and related tetranortriterpenoids.     

Stem respiratory potential in six softwood and four hardwood tree species in the central cascades of Oregon

Mature and old growth trees of varying sapwood thickness were compared with regard to stem respiration. An increment core-based, laboratory method under controlled temperature was used to measure tissue-level respiration (termed respiratory potential) of ten different tree species. Bark (dead outer and live inner combined), sapwood, and heartwood thickness measurements were used to predict sapwood volume from stem diameter (including bark) for four of the ten species. These predictions of sapwood volume were used to scale respiratory potential to the main-bole level (excluding all branches). On the core level, species that maintained narrow sapwood (8–16% of bole radius) such as Pseudotusga menziesii, Taxus brevifolia, and Thuja plicata, had sapwood respiratory potentials in the lower bole that were 50% higher (P<0.05) than species with wide sapwood (>16% of bole radius), such as Abies amabilis, Pinus monticola, and Tsuga heterophylla. This pattern was not observed for inner bark respiratory potential, or for sapwood respiratory potential within the crown. On the main-bole level, respiratory potential per unit volume was inversely correlated to the live bole volumetric fraction (inner bark plus sapwood divided by whole bole volume) (Adj. R²=0.6). Specifically, tree species with 18–20% of the main bole alive potentially respired 1.3–3 times more per unit live bole volume than species with over 40%, suggesting that the live bole was less metabolically active in tree species that maintained large volumes of sapwood.     

Can Norway spruce trees be ‘vaccinated’ against attack by Ips typographus?

1 A field experiment was carried out to test the hypothesis that treatment of Norway spruce trees with the Ips typographus-transmitted blue-stain fungus Ceratocystis polonica enhances tree resistance to later mass attack by this bark beetle. 2 Twenty-five mature trees were pretreated by inoculating a non-lethal dose of the fungus into the bark, while 18 trees served as untreated controls. Three and a half weeks after treatment a bark beetle attack was initiated by attaching dispensers with I. typographus pheromone to the tree trunks. 3 A significantly larger proportion (67%) of the control trees than of the pretreated trees (36%) were killed by the beetle attack. The result is discussed in relation to recent results regarding defence mechanisms in Norway spruce trees.     

Variations in bark-stripping by red deer Cervus elaphus across Europe

• 1 The literature on bark‐stripping by red deer Cervus elaphus in Europe is reviewed to reveal quantitative variation in this behaviour and relate it to deer density and local characteristics such as dominant tree species, occurrence of artificial feeding, altitude, region and size of the study site. We also review the importance of bark in red deer diets over the seasons and discuss the causes of bark‐stripping, focusing on the significance of bark as food.
• 2 Over the 36 sites examined, the rate of bark‐stripping was highly variable (from 0 to 84% of susceptible trees debarked), with less damage in Scotland than in other European sites for which bark‐stripping rates were higher at high red deer density. Altitude, the size of the study site, the number of dominant tree species and the occurrence of artificial feeding do not significantly relate to the rate of bark‐stripping.
• 3 Bark sometimes made up a large proportion of red deer diet (> 10%), especially in areas with severe winters (high levels of snow), whereas in study sites with mild winters, bark was practically not eaten at all.
• 4 These results suggest that severe bark‐stripping could be related to a reduction in food resource availability. This food availability hypothesis needs to be better documented, dealing particularly with the possible interaction between food availability and red deer density.

     

SITE-SPECIFIC CHANGES IN THE BARK TEXTURE OF QUERCUS FALCATA MICHX. (SOUTHERN RED OAK)

A method of measuring bark texture is described. This method is used to show that bark roughness in Quercus falcata Michx. (southern red oak) increases from mesic to xeric sites. This change does not appear to be explained by variability in tree size and age, or by changes in the relative abundances of varieties falcata and pagodaefolia. We suggest that it may be due to postulated relationships between bark texture (or the associated property of outer bark thickness) and tree fire resistance, nutrient conservation or susceptibility to pathogens.     

The bark of trees: thermal properties,microclimate and fauna

Summary The thermal properties of four different types of bark were investigated on twentyfour central European tree species using thermocouples. Tree species with white bark avoid overheating of their surface by reflection of the radiation. Species with fissured and scaly barks shade inner parts of their bark. Some tree species with fissured barks show high insulation across the bark. Smooth and thin barks show no adaptation to avoid overheating. These tree species (in central Europe e.g. Fagus sylvatica) have to form closed stands and are not able to occur in open stands as tree species with structured or white bark types.The arthropod fauna of the same bark types was studied on six tree species using three collecting methods. The microclimate on the bark determines the number per cm² of some species, some are active during winter, and other tend to aptery and reproduce parthenogenetically. On smooth bark one species dominates, whereas a highly diverse fauna lives on fissured barks. The occurrence of species on bark is determined by the microstructure, microclimate and consistence of the epiphyts. If tree species alter within forest ecosystems the dominant species on bark will persist. Specialists of fissured barks will die out if tree species with smooth bark (Fagus sylvatica) form the central European forest.     

Behavior of male and female spruce bark beetles,Ips typographus,on the bark of host trees during mass attack

The behavior of 118 spruce bark beetles, Ips typographus,was observed on trees under colonization. Most individuals were followed from when they landed until they entered or left the tree. Both males and females spent most time inspecting crevices and searching for a place to start boring or for a hole to enter. These behaviors accounted for 87 and 70% of all behavioral acts recorded for males and females, respectively. Females entered galleries with males only after a period of pushing at the gallery entrance. Males spent on average 3 min and females 4 min on the bark before entering or leaving the tree. Thirty-three percent of the beetles eventually entered the tree, 31% flew away, 35% dropped from the host, and one beetle was eaten by a predator. The results are discussed in relation to the question of mate choice in bark beetles and to studies on attack dynamics of spruce bark beetle populations.     

Tree mortality episodes in the intact Picea abies‐dominated taiga in the Arkhangelsk region of northern European Russia

Question: What were the temporal patterns and rates of tree mortality in a recent episodic tree mortality event? Have similar events occurred in the past, and does climatic variability play a role in the disturbance regime? Location: Intact Picea abies‐dominated taiga in the Arkhangelsk region, northwestern Russia. Methods: We reconstructed the past tree mortality and disturbance history by applying dendroecological methods in five forest stands and related these to climatic data. The role of other potential causes of tree mortality was assessed in a field inventory. Results: The recent episode lasted from 1999 to 2004, influenced all stands studied, and killed on average 21% of trees with a diameter of over 10 cm at 1.3‐m height. The annual tree mortality rate in the decades preceding this episode was 0.49%. During the past 200 years, the stands have experienced chronic small‐scale disturbances, with several irregular disturbances of moderate severity. The recent episode was associated with abundant signs of the bark beetle Ips typographus. Furthermore, the timing of both the recent tree mortality episode and the past disturbance events was associated with dry summers. Conclusion: The results indicate a connection between climatic variability and forest dynamics, the likely driving factors being droughts and bark beetles. In the context of the past 200 years, the recent episode was potentially at the higher end of the range of disturbance variability in terms of severity and spatial extent. This has ecological implications in a changing climate, potentially influencing ecosystem structure and long‐term dynamics.     

Conifer phenolic resistance markers are bark beetle antifeedant semiochemicals

1 Phenols are important in conifer resistance to fungi associated with bark beetles and as markers for resistance to beetle mass‐attacks. However, the mechanism of phenolic activity in conifer resistance to Ips typographus bark beetle remains unclear. 2 By a new bioassay, we tested the effect on host acceptance and tunnelling of male and female I. typographus in an artificial gallery (13 mm long) for 4 h (no‐choice test). To the artificial diet, an aliquot of host (catechin, taxifolin, or resveratrol) or nonhost (E‐conophthorin) compounds was added, singly tested at doses close to those of tree bark. Host acceptance and tunnelling was measured by the amount of diet removed by the insects. 3 All tested chemicals decreased the tunnelling activity of I. typographus, with an antifeedant effect stronger in males and increasing with dose. No mortality was recorded. The nonhost volatile spiroketal, E‐conophthorin, had the highest antifeedant activity both in males and females. Among host compounds, effects and dose–response were weak in females. Both catechin and E‐conophthorin gave a 50% reduction of tunnelling at a concentration of 0.1% for males, the pioneering (host selecting) sex in Ips. The threshold of activity for host compounds to males was at concentrations of 0.03–0.1%, which corresponds to, or is less than, the concentrations reported from spruce host bark. 4 The results allow us to support the hypothesis of a direct behavioural antifeedant mechanism for resistance from those phenols that are particularly active for the pioneering males during tree attack.     

Fire‐induced tree mortality in a neotropical forest: the roles of bark traits,tree size,wood density and fire behavior

Large‐scale wildfires are expected to accelerate forest dieback in Amazônia, but the fire vulnerability of tree species remains uncertain, in part due to the lack of studies relating fire‐induced mortality to both fire behavior and plant traits. To address this gap, we established two sets of experiments in southern Amazonia. First, we tested which bark traits best predict heat transfer rates (R) through bark during experimental bole heating. Second, using data from a large‐scale fire experiment, we tested the effects of tree wood density (WD), size, and estimated R (inverse of cambium insulation) on tree mortality after one to five fires. In the first experiment, bark thickness explained 82% of the variance in R, while the presence of water in the bark reduced the difference in temperature between the heat source and the vascular cambium, perhaps because of high latent heat of vaporization. This novel finding provides an important insight for improving mechanistic models of fire‐induced cambium damage from tropical to temperate regions. In the second experiment, tree mortality increased with increasing fire intensity (i.e. as indicated by bark char height on tree boles), which was higher along the forest edge, during the 2007 drought, and when the fire return interval was 3 years instead of one. Contrary to other tropical studies, the relationship between mortality and fire intensity was strongest in the year following the fires, but continued for 3 years afterwards. Tree mortality was low (≤20%) for thick‐barked individuals (≥18 mm) subjected to medium‐intensity fires, and significantly decreased as a function of increasing tree diameter, height and wood density. Hence, fire‐induced tree mortality was influenced not only by cambium insulation but also by other traits that reduce the indirect effects of fire. These results can be used to improve assessments of fire vulnerability of tropical forests.     

Pinus contorta var. yukonensis var. nov. (Pinaceae) from south–central Yukon,Canada

Pinus contorta Douglas ex Loudon var. yukonensis W. L Strong (Yukon pine), a new variety, is described from south–central Yukon, Canada. It is a stout tree with long lower branches, often with paired lower and forked upper stems, and weakly platy to smooth gray bark on the lower stem, when < 70 years old; with bark on the lower stem becoming furrowed and platy on older trees. This contrasts with the narrow columnar habit, simple stem, and scaly bark associated with the more common inland var. latifolia. In several respects, var. yukonensis is more similar to var. contorta (a coastal variant), although occupying a different ecological regime and spatially disjunct (i.e., a high latitude and high elevation variant). Variety yukonensis may represent the taxon alluded to in previous leaf oil terpene and genetic studies, and a possible Beringian survivor.     

Fungi Vectored by the Bark Beetle Ips typographus Following Hibernation Under the Bark of Standing Trees and in the Forest Litter

The bark beetle Ips typographus has different hibernation environments, under the bark of standing trees or in the forest litter, which is likely to affect the beetle-associated fungal flora. We isolated fungi from beetles, standing I. typographus-attacked trees, and forest litter below the attacked trees. Fungal identification was done using cultural and molecular methods. The results of the two methods in detecting fungal species were compared. Fungal communities associated with I. typographus differed considerably depending on the hibernation environment. In addition to seven taxa of known ophiostomoid I. typographus-associated fungi, we detected 18 ascomycetes and anamorphic fungi, five wood-decaying basidomycetes, 11 yeasts, and four zygomycetes. Of those, 14 fungal taxa were detected exclusively from beetles that hibernated under bark, and six taxa were detected exclusively from beetles hibernating in forest litter. The spruce pathogen, Ceratocystis polonica, was detected occasionally in bark, while another spruce pathogen, Grosmannia europhioides, was detected more often from beetles hibernating under the bark as compared to litter. The identification method had a significant impact on which taxa were detected. Rapidly growing fungal taxa, e.g. Penicillium, Trichoderma, and Ophiostoma, dominated pure culture isolations; while yeasts dominated the communities detected using molecular methods. The study also demonstrated low frequencies of tree pathogenic fungi carried by I. typographus during its outbreaks and that the beetle does not require them to successfully attack and kill trees.     

Attack preference of Ips pini on Pinus ponderosa in northern Arizona: tree size and bole position

1 The present study investigated the influence of tree size and bole position on selection of ponderosa pines (Pinus ponderosa Dougl. ex Laws.) by Ips pini Say (Coleoptera: Curculionidae: Scolytinae) in a northern Arizona forest. Ips pini were attracted to lower and upper bole positions with pheromone lures, and the number of attacks counted. In addition, tree phloem and bark thickness, resin flow in response to wounding and characteristics of tree water and carbon relations were all measured. 2 Bark thickness was the most strongly related tree characteristic to number of I. pini attacks. Thin bark and a high number of attacks occurred at the lower bole position of 10‐cm diameter trees and upper bole positions of 23‐, 35‐, and 50‐cm diameter trees. 3 Phloem thickness increased directly with tree size, was greater at the upper bole compared with the lower bole, but was not associated with number of attacks. 4 Resin flow did not differ strongly over tree sizes or bole positions, and was not related to number of I. pini attacks. Attacks were not related to characteristics of tree carbon and water relations measured during I. pini flights.