Suitability of native and ornamental oak species in California for Agrilus auroguttatus

Goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), is a new invasive species in southern California, USA. The extent of the host range of this insect is not known, but this knowledge will have a major impact on assessment of the risks that this pest poses to oaks [Quercus spp. (Fagaceae)]. We conducted laboratory tests to determine the potential suitability of native and ornamental oak species for larvae and adults of A. auroguttatus. We infested 179 cut logs (from 163 different trees) with eggs or larvae, measured neonate survival and, after 5 months, counted feeding galleries, and noted the proportion of galleries with late instars. Initial larval survival was generally high when larvae penetrated the phloem (range: 62–98% among oak species), and low by the time larvae began to feed at the phloem/xylem interface (range: 0–25% among oak species). The majority of larvae that established a visible feeding gallery survived to the fourth instar (total of 73% for all oak species). Larval galleries were established with greater frequency in red oaks (Section Lobatae) compared with other oaks (19 vs. 7 or 4%). All red oaks tested (Q. agrifolia Née, Q. kelloggii Newberry, and Q. wislizeni A. DC.) were likely suitable hosts for larvae. Larvae were apparently able to feed on some of the other oaks (Q. chrysolepis Leibmann, Q. suber L., Q. lobata Née, and Q. douglasii Hook & Arn), although it remains unclear whether these species would be preferred hosts under natural conditions. Adult longevity and fecundity varied little by species of oak foliage fed to adults. The host range of A. auroguttatus is likely limited by suitability of oak species for the larval rather than the adult life stage. Results support published field observations that red oaks are more suitable hosts than white oaks.     

Utilization of evergreen and decidous oaks by the Californian oak moth Phryganidia californica

Summary The deciduous oak Quercus lobata was a better quality food source than the evergreen oak Q. agrifolia for the California oak moth Phryganidia californica: it supported higher growth rates, produced higher fecundity and survivorship, and higher efficiency of conversion of ingested food (ECI), efficiency of conversion of digested food (ECD) and efficiency of nitrogen utilization (NUE). Total polyphenols and astringency were similar in both oak speices, whereas water content was lower and leaf specific weight higher in Q agrifolia than in Q lobata. Nitrogen was higher in new leaves of Q agrifolia but higher in mature leaves of Q. lobata. Contrary to most other studies, total polyphenols and astringency were higher in new than in mature leaves. These results confirmed my hypothesis that the differences in leaf quality associated with the evergreen or deciduous life form would translate into food of differing quality for herbivores.     

Goldspotted oak borer effects on tree health and colonization patterns at six newly‐established sites

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Adaptation to host plants may prevent rapid insect responses to climate change

We must consider the role of multitrophic interactions when examining species' responses to climate change. Many plant species, particularly trees, are limited in their ability to shift their geographic ranges quickly under climate change. Consequently, for herbivorous insects, geographic mosaics of host plant specialization could prohibit range shifts and adaptation when insects become separated from suitable host plants. In this study, we examined larval growth and survival of an oak specialist butterfly (Erynnis propertius) on different oaks (Quercus spp.) that occur across its range to determine if individuals can switch host plants if they move into new areas under climate change. Individuals from Oregon and northern California, USA that feed on Q. garryana and Q. kelloggii in the field experienced increased mortality on Q. agrifolia, a southern species with low nutrient content. In contrast, populations from southern California that normally feed on Q. agrifolia performed well on Q. agrifolia and Q. garryana and poorly on the northern, high elevation Q. kelloggii. Therefore, colonization of southern E. propertius in higher elevations and some northern locales may be prohibited under climate change but latitudinal shifts to Q. garryana may be possible. Where shifts are precluded due to maladaptation to hosts, populations may not accrue warm‐adapted genotypes. Our study suggests that, when interacting species experience asynchronous range shifts, historical local adaptation may preclude populations from colonizing new locales under climate change.     

Adaptation to oak and other fibrous,phenolic-rich foliage by a small mammal,Neotoma fuscipes

Summary Neotoma fuscipes, a small mammalian herbivore with apparently generalized food habits, was laboratory tested to determine its degree of dietary specialization. Woodrats from both oak woodland and coastal sage communities preferred Quercus agrifolia leaves (containing 40% phenolics and about 16% condensed tannin) over foliage from other dominant species. Approximately one-third of the oak phenolics and less than 10% of the oak condensed tannin remained in the feces. Their performance on pure oak leaves was comparable to that on a mixed diet of Quercus, Salvia, Eriogonum, and Rhus, with respect to weight maintenance, digestive efficiency and total amount ingested. Digestive efficiency was low on the oak diet (55%) relative to Salvia (77%), and to achieve similar weight levels, approximately twice as much oak as Salvia was ingested. Woodrats retained more nitrogen as oak consumption increased. Intake of oak and other foods increased with each experimental day. A sympatric species, N. lepida, was unable to maintain weight on oak leaves, although its digestive and polyphenolic-degrading capabilities, and nitrogen retention efficiency were equivalent to those of N. fuscipes. On a weight-adjusted basis, N. lepida ate about half as much oak per day as N. fuscipes. Oak intake may have been reduced by an inability to rapidly degrade fiber, which constitutes about 30% of the oak diet. In natural populations, N. fuscipes selectively feeds on evergreen sclerophyll vegetation high in fiber, tannins and related polyphenolics. Individuals ingest 2–3 plant types at a time, with a single species (oak when available) constituting most of the material consumed. Neotoma lepida diets are also dominated by a single species. The diversity of plant types eaten by different populations of N. lepida suggests that local dietary specializations may be developmentally acquired.     

Seasonal variations of gas exchange, photosynthetic pigments, and antioxidants in Turkey oak (Quercus cerris L.) and Hungarian oak (Quercus frainetto Ten.) of different age

Seasonal changes in physiological and biochemical parameters were studied in 35-, 55- and 140-year-old trees of Turkey oak (Quercus cerris L.) and Hungarian oak (Q. frainetto Ten.), growing in natural stands in Eastern Balkan Mountains (Bulgaria). During the seasonal drought period (August), assimilation activity, transpiration rate, stomatal conductance and water potential had a seasonal minimum in all the studied tree ages and species. The foliar concentrations of glutathione, ascorbate, α-tocopherol, as well as photosynthetic pigments in oak leaves were significantly affected by season. With the increasing age of the studied trees, we observed a decrease of the physiological activity and an increase of the antioxidants’ accumulation. Both the species were drought tolerant and anisohydric, where Q. frainetto exhibited higher rates of gas exchange than Q. cerris. Moreover, they differed in the extent of increase in the foliar antioxidants and carotenoids.     

Seasonal variation in leaf chemistry of the coast live oak Quercus agrifolia and implications for the California oak moth Phryganidia californica

Summary The perennial foliage of the California coast live oak (Quercus agrifolia Nee) permits herbivores to feed on this oak species throughout the year. Patterns of herbivory for a two-year period on Q. agrifolia were observed in relation to seasonal and age-related changes in the nutritional and defensive characteristics of leaves. Nitrogen and phosphorus contents were higher in new leaves compared to mature foliage. Structural compounds (e.g., cellulose) in leaves rapidly increased with age. Concentrations of tatal phenolics (Folin-Denis) and astringency were higher in new foliage, and concentrations of condensed tannins gradually increased as the leaves matured. Peaks of herbivore damage were observed in June and in September–October, and were caused by outbreaks of the California oak moth (Phryganidia californica). P. californica, a bivoltine oak specialist, exhibited feeding preferences in June for old leaves over emerging leaves, and showed no preferences for leaf classes in September. These results suggest that P. californica is adapted to survive on nutritionally poor foliage and to circumvent quantitative defenses such as condensed tannins.     

Long‐term and interactive effects of different mammalian consumers on growth,survival, and recruitment of dominant tree species

Throughout the world, numerous tree species are reported to be in decline, either due to increased mortality of established trees or reduced recruitment. The situation appears especially acute for oaks, which are dominant features of many landscapes in the northern hemisphere. Although numerous factors have been hypothesized to explain reductions in tree performance, vertebrate herbivores and granivores may serve as important drivers of these changes. Here, using data from 8‐ and 14‐year‐old exclosure experiments, we evaluated the individual and interactive effects of large and small mammalian herbivores on the performance of three widespread oak species in California—coast live oak (Quercus agrifolia), California black oak (Q. kelloggii), and Oregon white oak (Q. garryana). Although impacts varied somewhat by species and experiment, herbivory by black‐tailed deer (Odocoileus hemionus columbianus) reduced the height and survival of juvenile coast live oaks and altered their architecture, as well as reduced the abundance of black oak seedlings, the richness of woody species and the cover of nonoak woody species. Small mammals (Microtus californicus and Peromyscus maniculatus) had even more widespread effects, reducing the abundance of black oak seedlings and the height and cover of all three oak species. We also detected numerous interactions between small mammals and deer, with one herbivore having positive or negative effects on oak abundance and cover when the other herbivore was either present or absent. For example, deer often had negative effects on seedling abundance only when, or even more so when, small mammals were present. In summary, mammalian consumers play crucial roles in limiting oak recruitment by reducing seedling abundance, maintaining trees in stunted states, and preventing them from reaching sapling stages and becoming reproductive. Interactions between large and small mammals can also alter the intensity and direction of their effects on trees.     

Distinct male reproductive strategies in two closely related oak species

Reproductive strategies of closely related species distributed along successional gradients should differ as a consequence of the trade‐off between competition and colonization abilities. We compared male reproductive strategies of Quercus robur and Q. petraea, two partly interfertile European oak species with different successional status. In the studied even‐aged stand, trees of the late‐successional species (Q. petraea) grew faster and suffered less from intertree competition than trees of the early‐successional species (Q. robur). A large‐scale paternity study and a spatially explicit individual‐based mating model were used to estimate parameters of pollen production and dispersal as well as sexual barriers between species. Male fecundity was found to be dependent both on a tree's circumference and on its environment, particularly so for Q. petraea. Pollen dispersal was greater and more isotropic in Q. robur than in Q. petraea. Premating barriers to hybridization were strong in both species, but more so in Q. petraea than in Q. robur. Hence, predictions based on the competition–colonization trade‐off are well supported, whereas the sexual barriers themselves seem to be shaped by colonization dynamics.     

Environmental suitability for Agrilus auroguttatus (Coleoptera: Buprestidae) in Mexico using MaxEnt and database records of four Quercus (Fagaceae) species

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Acorns containing deeper plumule survive better: how white oaks counter embryo excision by rodents

Several squirrel species excise the embryo of acorns of most white oak species to arrest germination for long‐term storage. However, it is not clear how these acorns counter embryo excision and survive in the arms race of coevolution. In this study, we simulated the embryo excision behavior of squirrels by removing 4 mm of cotyledon from the apical end of white oak acorns differing in embryo depths to investigate the effects of embryo excision on acorn germination and seedling performance of white oak species. The embryo depth in the cotyledons was significantly different among white oak acorns, with Quercus mongolica containing the embryo most deeply in the acorns. We found that artificial embryo excision significantly decreased acorn germination rates of Quercus variabilis, Quercus acutissima, Quercus aliena, Quercus aliena var. acutiserrata, Quercus serrata. var. brevipetiolata but not Q. mongolica. Artificial embryo excision exerted significant negative impacts on seedling performance of all oak species except Quercus aliena. Our study demonstrates the role of embryo depth of acorns in countering embryo excision by squirrels and may explain the fact that squirrels do not perform embryo excision in acorns of Q. mongolica with deeper embryos. This apparent adaptation of acorns sheds light on the coevolutionary dynamics between oaks and their seed predators.     

Flooding effects on starch partitioning during early growth of two oak species

Pedunculate (Quercus robur L.) and sessile (Q. petraea [Matt.] Liebl.) oaks are the most common oak species in Western Europe. They are known to display different ecological requirements, particularly relative to root hypoxia induced by flooding: In a glasshouse study of seedlings, we quantified the effects of flooding on starch mobilization from cotyledons and starch partitioning. Growth and distribution of lateral roots were also measured. The above-ground growth of Q. robur was less affected by flooding than that of Q. petraea which failed to develop a second flush. Root growth was also severely inhibited, particularly in Q. petraea. In Q. robur, lateral root initiation as well as elongation was restricted to the soil surface layer. Flooding markedly reduced total growth and concentrations of in all components except stems. Starch mobilization from cotyledons was delayed by flooding, especially in Q. robur seedlings. Under flooding, the decrease of cotyledons dry mass and starch content in Q. robur was lower than in Q. petraea, whereas Q. robur displayed larger growth than Q. petraea. The features of carbohydrate management may be crucial in the observed differences in flooding tolerance of these species.     

Natural hybridisation among Quercus glabrescens,Q. rugosa and Q. obtusata (Fagaceae): Microsatellites and secondary metabolites markers

• Natural hybridisation has significant ecological, genetic and evolutionary consequences altering morphological and chemical characters of individuals. Quercus glabrescens, Q. rugosa and Q. obtusata are white oak species well separated by their morphological characters when they occur in allopatry in Mexican temperate forests. However, in sympatry, individuals with atypical morphology have been observed, suggesting hybridisation events.
• In this study, we determined, with microsatellites and secondary metabolites, if interspecific gene flow occurs when these three oak species coexist in sympatry. In total, 180 individuals belonging to seven populations [three allopatric (one for each parental species) and four sympatric sites] were analysed.
• Allopatric populations represent well‐defined genetic groups and the sympatric populations showed genetic evidence of hybridisation between Q. glabrescens × Q. rugosa and Q. glabrescens × Q. obtusata. The hybridisation percentage varied between sites and combination of involved species. We registered the presence of unique flavonoid compounds for Q. glabrescens (caffeic acid and flavonol 2), Q. rugosa (flavonol 5) and Q. obtusata (flavonol 1). Three compounds (quercetin rhamnoside, flavonol 3 and alkyl coumarate) were expressed in all taxa. Finally, the hybrid genotypes identified in this study (Q. glabrescens × Q. rugosa and Q. glabrescens × Q. obtusata) showed specific chemical profiles, resulting from a combination of those of their parental species.
• These results show that hybridisation events between these oak species alter chemical expression of secondary metabolites, creating a mosaic of resources and conditions that provide the substrate for different combinations of foliar‐associated species such as herbivores, endophytic fungi or epiphyte plants.

     

Radial growth and climate responses of white oak (Quercus alba) and northern red oak (Quercus rubra) at the northern distribution limit of white oak in Quebec, Canada

Aim The objectives of this study were: (1) to compare radial growth patterns between white oak (Quercus alba L.) and northern red oak (Quercus rubra L.) growing at the northern distribution limit of white oak; and (2) to assess if the radial growth of white oak at its northern distribution limit is controlled by cold temperature. Location The study was conducted in three regions of the Ottawa valley in southern Québec. All stands selected were located at the northern limit of distribution of Q. alba. Methods Twelve mixed red and white oak stands were sampled and increment cores were extracted for radial growth analyses. For each oak species, 12 chronologies were derived from tree‐ring measurement (residual chronologies). Principal components analysis and redundancy analysis were used to highlight the difference between radial growth in both species and to determine their radial growth–climate association. Results There was little difference between the radial growth of each species; Q. alba, however, exhibits more year‐to‐year variation in growth than Q. rubra. More than 65% of the variance in radial growth was shared among sites and species. Both species showed a similar response to climate, which suggested that the limit of distribution of Q. alba might not be determined by effects on growth. Both species had a classic response to climate and drought in the early growing season. Main conclusions The northern distribution limit of Q. alba does not appear to be directly controlled by effects on growth processes as indicated by the similarities in radial growth and response to climate between the two species. The location of the stands on southern aspects suggested that cold temperature could have been a major factor controlling the distribution limit of Q. alba. However, it is speculated that stands growing on southern aspects may be more prone to forest fires or to drought, which would favour the maintenance and establishment of oaks, and of Q. alba in particular. Models relating the northern distribution limits of species to broad climate parameters like annual mean temperature will need to be reviewed to incorporate more biologically relevant information. Such assessments will in turn provide better estimates of the effect of climate changes on species distribution.     

Nitrogen sink strength of ectomycorrhizal morphotypes of <Emphasis Type="Italic">Quercus douglasii</Emphasis>, <Emphasis Type="Italic">Q. garryana</Emphasis>, and <Emphasis Type="Italic">Q. agrifolia</Emphasis> seedlings grown in a northern California oak woodland

Little information is known on what the magnitude of nitrogen (N) processed by ectomycorrhizal (ECM) fungal species in the field. In a common garden experiment performed in a northern California oak woodland, we investigated transfer of nitrogen applied as ¹⁵NH₄ or ¹⁵NO₃ from leaves to ectomycorrhizal roots of three oak species, Quercus agrifolia, Q. douglasii, and Q. garryana. Oak seedlings formed five common ectomycorrhizal morphotypes on root tips. Mycorrhizal tips were more enriched in ¹⁵N than fine roots. N transfer was greater to the less common morphotypes than to the more common types. ¹⁵N transfer from leaves to roots was greater when , not , was supplied. ¹⁵N transfer to roots was greater in seedlings of Q. agrifolia than in Q. douglasii and Q. garryana. Differential N transfer to ectomycorrhizal root tips suggests that ectomycorrhizal morphotypes can influence flows of N from leaves to roots and that mycorrhizal diversity may influence the total N requirement of plants.     

Phytophthora quercetorum sp. nov., a novel species isolated from eastern and north-central USA oak forest soils

Isolates belonging to an undescribed Phytophthora species were frequently recovered during an oak forest soil survey of Phytophthora species in eastern and north-central USA in 2004. The species was isolated using an oak leaf baiting method from rhizosphere soil samples collected from Quercus rubra, Q. macrocarpa, and Q. phellos. This species is formally described as P. quercetorum. It is homothallic and has aplerotic oogonia and paragynous antheridia. It produces papillate sporangia (occasionally bipapillate) of ovoid-elongated shapes. Its temperature optimum for growth is ca 22.5 °C with the upper limit of ca 32.5 °C. P. quercetorum differs from the morphologically related P. quercina in producing distinct submerged colony-patterns, different growth-temperature requirements, and oogonial shapes and sizes. Phylogenetic analyses using seven nuclear loci supported P. quercetorum as a novel species within clade 4, closely related to P. arecae, P. palmivora, P. megakarya, and P. quercina.     

Vegetation associations of birds wintering in a California oak woodland

Conditions in wintering areas can have persistent effects throughout the year for bird populations. Most studies, however, are conducted during the breeding season and the non‐breeding ecology of many species remains poorly understood. We assessed vegetation associations of birds wintering in a diverse, well‐structured oak woodland in coastal‐central California. We calculated density estimates for 20 bird species and examined correlations between bird densities and vegetation characteristics as described by a principal components analysis. Ruby‐crowned Kinglets (Regulus calendula) and Hutton's Vireos (Vireo huttoni) were positively associated with vegetation characteristic of dense, floristically diverse woodland with abundant coast live oak (Quercus agrifolia) cover and a structurally complex understory. Conversely, White‐breasted Nuthatches (Sitta carolinensis) and Western Bluebirds (Sialia mexicana) were negatively correlated with dense woodland and occurred at the highest densities on plots with large widely spaced blue oaks (Q. douglasii) and a sparse understory. Thirteen of the remaining 16 species were more abundant in dense woodland patches, but these relationships were not statistically significant. Our results are in agreement with those of previous studies and suggest that evergreen oak trees and a dense understory provide key resources (e.g., cover and food) for some bird species during the winter. However, areas with large widely spaced deciduous oaks are important habitats for other species. We therefore suggest that dense patches of live oaks and shrubs should be retained, but a mosaic of dense and sparse woodland is necessary to accommodate the entire suite of oak‐woodland birds.     

Rodent acorn selection in a Mediterranean oak landscape

Quercus suber, Quercus ilex and Quercus coccifera (Cork, Holm and Kermes oaks, respectively) are common evergreen oak species that coexist in the landscapes of the western part of the Mediterranean basin. Rodents are the main acorn predators and thus one of the main factors for understanding recruitment patterns in oaks. In this paper we analyse to what extent mice prefer acorns from one oak species over another in three oak species studied using acorn removal experiments and video tape recordings. Twenty labelled acorns from each of the three Quercus species (60 acorns) were placed in 40 cm×40 cm quadrats on each plot. Because selection might vary as a result of the vegetation context, we performed the trials in the five main vegetation types within the study area (four replicates in each vegetation type) in order to control for habitat influences on rodent acorn preferences (a total of 20 plots). The removal of 1,200 acorns occurred within 68 days. Mice removed 98.7% of the acorns. Q. ilex acorns were preferred over Q. suber and Q. coccifera in all vegetation types except in pine forest, where no acorn preferences were detected. Acorn removal rates differed with vegetation type, correlating positively with shrub cover. The distance at which acorns were displaced by rodents (mean =4.6 m±5.1 SD) did not differ between acorn species, but varied among vegetation types. Bigger acorns of Q. coccifera were selected only after Q. ilex and Q. suber acorns were depleted, while no size selection was detected for the latter two species. Thus, we conclude that rodents show preference for some oak acorns and that landscape context contributes significantly to rodent activities and decisions.     

Ectomycorrhizal fungal communities of pedunculate and sessile oak seedlings from bare-root forest nurseries

In this study, we present the detailed molecular investigation of the ectomycorrhizal (ECM) community of Quercus petraea and Quercus robur seedlings grown in bare-root forest nurseries. In all tested oak samples, mycorrhizal colonization was nearly 100%. Morphological observation and molecular investigations (sequencing of fungal ITS rDNA) revealed a total of 23 mycorrhizal taxa. The most frequent and abundant fungal taxa were Hebeloma sacchariolens, Tuber sp., and Peziza sp.; from the detected fungal taxa, 20 were noted for Q. petraea and 23 for Q. robur. Depending on the nursery, the species richness of identified ECM fungal taxa for both oak species ranged from six to 11 taxa. The mean species richness for all nurseries was 5.36 and 5.82 taxa per Q. petraea and Q. robur sample, respectively. According to the analysis of similarity, ECM fungal communities were similar for Q. petraea and Q. robur (R = 0.019; p = 0.151). On the other hand, detected fungal communities were significantly different between nurseries (R = 0.927; p < 0.0001). Using the Spearman rank correlation, it was determined that the ectomycorrhizal diversity (in terms of richness, the Shannon diversity, evenness, and Simpson dominance indices) is significantly related to the soil parameters of each nursery. We conclude that individual nursery may be considered as separate ecological niches that strongly discriminate diversity of ECM fungi.     

The feeding behaviour of a sit-and wait-predator,Ranatra dispar (Heteroptera: Nepidae): optimal foraging and feeding dynamics

Summary Nitrogen and phosphorus flow in litterfall and throughfall were studied in two California Quercus species (the evergreen Q. agrifolia and deciduous Q. lobata) before, during, and after an outbreak of the California oak moth, Phryganidia californica. All of the foliage of both oak species was removed by the herbivore during the course of this outbreak. During the outbreak, total N and P flow to the ground more than doubled from Q. agrifolia and increased to a lesser extent from Q. lobata over the previous year. The composition of the litter during the outbreak year shifted so that in Q. agrifolia, almost 70% of the total N and P flow to the ground moved through frass and insect remains, while in Q. lobata, approximately 60% of the N and 40% of the P moved through frass and insect remains. Short-term leaching experiments showed that nitrogen was far more rapidly lost from Phryganidia frass than from leaf litter of either species. These results and the relative frequency of Phryganidia outbreaks suggest that this herbivore has significant effects on the nutrient cycling beneath these trees.