Think globally,manage locally: The importance of steady-state forest features for a declining songbird

Changes in historical forest composition and structure may have cascading effects throughout the forest community. Perhaps nowhere is there a better example of current forests that carry a legacy from their past than in eastern North America. The Cerulean Warbler (Dendroica cerulea), a declining Neotropical migratory bird of high conservation concern, is one excellent example of a species that seems to be sensitive to both landscape configuration and subtle features of eastern forests of North America. We used the Cerulean Warbler as a model species to demonstrate how an appreciation of fine-scale structural attributes of forests may improve our ability to conserve late-successional forest species. To do this we evaluated the extent to which multiscale habitat features were associated with density, spatial distribution, and nesting success of Cerulean Warblers in 12 mature forest sites in southeast Ohio, 2004–2006. Results suggest that adjacency of regenerating clear-cuts did not influence density or nesting success of Cerulean Warblers in adjacent mature forest. Instead, variation in demographic parameters was best explained by local habitat features. Density and nesting success were positively associated with canopy openness, numbers of large-diameter trees, and number of grapevines—all of which are typical of heterogeneous steady-state phase forests. Thus, improved management for Cerulean Warblers may require creating features (e.g., large canopy gaps) that mimic old-growth forests. Although fragmentation and habitat loss remain important contributors to population declines of many mature forest species, our work provides evidence that subtle changes in forest structure, particularly to features associated with old forests, warrant additional attention from the conservation community.     

Change in oak abundance in the eastern United States from 1980 to 2008

Although oaks (Quercus spp.) have historically dominated much of the forest land in eastern North America, a great deal of fragmentary and sometimes anecdotal evidence suggests that they have been yielding dominance in recent decades to other, typically more shade-tolerant species. Using FIA data, our work formally quantifies the change in oak abundance in the eastern U.S. during the period of 1980-2008. The results indicate that most areas in the eastern U.S. experienced some decline in oak abundance, but the decrease was not universal either geographically or among species. Declines were especially marked in the Central Hardwood Region, which lost oak abundance on 81% its forested area as measured by importance value (IV). Areas with a high oak abundance were more likely to see a reduction in abundance. Among all 25 species analyzed, eight species decreased significantly in IV while two increased. Both the top two most prevalent white oak species (white oak (Quercus alba) and post oak (Quercus stellata)) and red oak species (northern red oak (Quercus rubra) and black oak (Quercus velutina)) had significant decreases in density and IV. Water oak (Quercus nigra) is one of the red oak species that had a near universal increase of its abundance throughout its native range (83% of area). This study provided a comprehensive quantification of the dynamic of oak species in a regional-wide geographic context, which will provoke forest researchers and managers to revisit the oak decline problem by using knowledge from other regions and other species.     

Establishment and growth of oak (Quercus alba, Quercus prinus) seedlings in burned and fire-excluded upland forests on the Cumberland Plateau

Recurrent problems with regeneration of oaks (Quercus spp.) have been documented across a wide range of ecosystems. In oak-dominated forests of the central and Appalachian hardwood regions of the United States, a lack of competitive oak regeneration has been tied, in part, to fire suppression in these landscapes, and managers throughout the region are using prescribed fire to address this concern. To examine fire effects on oak regeneration, researchers have generally relied on inventories or population studies of existing seedlings. These studies are valuable but do not permit examination of the role of fire in enhancing the establishment and growth of new oak seedlings stemming from oak mast events. In this study, white (Quercus alba) and chestnut oak (Quercus prinus) acorn mast crops serendipitously occurred in year three (fall 2005) of a landscape-scale prescribed fire experiment. We examined establishment, survival, height and diameter of new seedlings on sites on the Cumberland Plateau in eastern Kentucky. Treatments were fire exclusion, a single prescribed fire (1x-burn; 2003), and repeated prescribed fire (3x-burn; 2003, 2004, and after acorn drop in 2006), all conducted in late spring. Initial densities of newly established chestnut and white oak seedlings were statistically similar across treatments (P = 0.42), despite fires on the 3x-burn site having occurred after acorns were on the ground. Oak seedling density was significantly predicted by oak basal area on all sites (R² = 0.12–0.46), except for chestnut oak on fire-excluded sites (R² = 0.04). Litter depth was less on 3x-burn sites compared to 1x-burn and fire-excluded sites, whereas canopy openness was greater on both burn treatments compared to fire-excluded sites. Seedling mortality was generally higher on fire-excluded sites compared to burn sites, especially for white oak. Oak seedling mortality in the first two growing seasons was significantly predicted by initial litter depth and open sky, with greater litter depth and lower percent open sky leading to higher mortality. In the third growing season none of the measured variables predicted chestnut oak seedling survival; for white oak, percent open sky remained a significant predictor of mortality. Initially, seedlings on the fire-excluded sites had similar height but smaller diameter; after three growing seasons there were few differences in seedling height or diameter among treatments. Our findings suggest a potential role for prescribed fire in establishing forest floor and light conditions that may enhance the success of new oak germinants, although different responses among species may suggest the need to target management for individual oak species.     

Silvicultural methods of oak regeneration with special respect to shade tolerant mixed species

An overview is presented of the silviculture of pedunculate oak (Quercus robur L.) and sessile oak (Q. petraea Liebl.) in Germany. This presentation is confined to less dry to moist and stagnic gleysol sites, where the intention is to produce primary timbers in long rotation periods. Incorporation of a shade tolerant species as an admixture species is indispensable to the suppression of epicormic branches. The most frequent and recommended admixture species for this purpose is beech (Fagus sylvatica L.). Because beech is competitively stronger than the oaks on these sites, silvicultural measures must be taken to keep it in check. According to the classic silviculture which has taken shape in the Spessart and Pfälzer forests, the desired goal of a two-storeyed stand with oak in the overstorey and beech in the understorey is achieved by means of heavy seeding beneath an open and rapidly cleared canopy. Because this procedure resembles clear cutting and entails its recognized disadvantages, trials have recently been undertaken to regenerate oaks in a silviculture with permanent canopy cover. Trials with young oaks show that they still achieve satisfactory growth at 15–20% of full light. Accordingly, successful regeneration is possible in beech stands under an open canopy or in gaps, and under an approximately closed canopy cover in pine stands due to their more penetrable crowns. The problematic aspect of this, especially in beech stands, is that young beeches become competitively stronger than oaks as canopy cover increases. This requires great effort in restraining the beech during cleanings and thinnings. Furthermore, browsing by wild animals must be minimized as it exclusively affects oak. As a result, however, there will be significantly fewer oaks and more beeches in the dominant layer than when using classic methods.     

Identification and distribution of Armillaria species associated with an oak decline event in the Arkansas Ozarks

Forests in the Ozark Mountains of northern Arkansas recently experienced a widespread oak decline event. Armillaria, a root rot fungus, has been associated with other oak decline events and may have been an important contributing factor to tree mortality in this event. Although Armillaria has been identified from the Ozark Mountains in Missouri, it has never been investigated in the Arkansas Ozarks. Molecular diagnostic techniques were used in this study to identify species of Armillaria present on roots removed from dead trees of two common oak species, northern red oak, Quercus rubra L., and white oak, Q. alba L., from three geographic areas and on three topographic positions – ridges, south‐ and west‐facing benches. Armillaria(A. mellea, A. gallica or A. tabescens) was identified from 31% of root samples taken from 102 trees in seven of nine sample plots. Armillaria mellea, occurred most often (20 samples, both oak species on seven plots) followed by A. gallica (10 samples, northern red oak only on four plots), and A. tabescens occurred twice (on northern red oak in a single plot). Thus, all three Armillaria species occurred on northern red oaks while A. mellea was the only species recovered from white oaks. Results varied by topographic position with samples from tree roots on ridges having the fewest positive identifications, one of 29. West‐facing benches had the highest positive samples with 20 of 41 testing positive and trees on south‐facing benches were intermediate with 11 of 32 samples from infected trees. This study documents the occurrence of three species of Armillaria in the Arkansas Ozarks and their association with oak mortality resulting from an oak decline event coupled with a red oak borer, Enaphalodes rufulus, outbreak. Further, it documents some potential variation in host/pathogen combinations and forest site conditions.     

The burning characteristics of southeastern oaks: Discriminating fire facilitators from fire impeders

In southeastern pine–oak ecosystems, ecological restoration targets oaks for removal by chemical, mechanical, burning, or combinations of treatments. Managers often pursue oak removal indiscriminately despite the poorly understood historical structure, cover, and ecological function within these ecosystems. Restoration treatments often cite the impediment that oak litter represents to prescribed fire spread and effectiveness. We evaluated the burning characteristics of eight southeastern Quercus spp. by burning collected litter under controlled conditions in a combustion chamber. Replicated burns consisted of 15 g of litter on a 35 cm × 35 cm grid of xylene-soaked cotton strings. Burning characteristics measured included maximum flame height (cm), flaming duration (s), smoldering duration (s), residual ash, and mass loss rate (g s⁻¹). We compared all 8 oaks using ANCOVA, with litterbed depth as a covariate. The oaks differed for all burning characteristics measured (P < 0.001). Rank comparisons placed Quercus stellata and Quercus laevis as the species with greatest fire intensity, sustainability, and consumability, equivalent in many measures to longleaf pine and other fire resisters. Quercus virginiana and Quercus hemisphaerica burned with the least intensity, sustainability, and consumability, burning similarly to sand pine and other fire evaders. These results show that oaks common to southeastern United States ecosystems have litter properties, similar to pines, which vary in their ability to sustain fire. Understanding the pyric properties of oak species also suggests that managers prioritize removal of species that hinder prescribed fire effectiveness for restoration of southeastern USA pine–oak ecosystems.     

How we improved a landscape study of species richness of beetles in woodland key habitats, and how model output can be improved

In an earlier study (Franc et al., 2007), local species richness of saproxylic oak beetles (including red-listed beetles) in forests was predicted mainly by the landscape (area of woodland key habitat within 1 km of plots). Such results are important for conservation work, but need to be backed up well, for reliable advice. We tested a two-stage method that improved our earlier models and our advice for conservation planning. We studied temperate mixed forest, rich in oaks Quercus robur/Quercus petraea, in a large landscape in Sweden. Franc et al. (2007) analysed 21 forests. Here we selected the significant explanatory variables (predictors) and other biologically relevant predictors, used the earlier 21 forests and sampled 11 new forests such that we expanded the range on the axes of the predictors. We collected in total 320 species of saproxylic oak beetles (23,137 individuals) of which 65 and 38 were red-listed (IUCN criteria, Swedish list 2000 and 2005, respectively). We partly confirmed our original results, but the results also changed in important ways: local species richness is now predicted by a combination of local, landscape and regional factors. Moreover, a local variable (dead wood) was the main predictor of saproxylic oak beetles (all species included), while for red-listed saproxylic oak beetles the landscape (woodland key habitat within 1 km of plots) was the main predictor, of local species richness. Thus, species richness of red-listed saproxylic oak beetles seems to depend mainly on landscapes factors, while total species richness of saproxylic oak beetles seems to depend more on local stand factors. We conclude that a two-stage research design can be useful in landscape and conservation studies, especially for species-rich taxa that require large samples per site.     

Stand and individual tree characteristics associated with Enaphalodes rufulus (Haldeman) (Coleoptera: Cerambycidae) infestations within the Ozark and Ouachita National Forests

Many oak decline events have been reported within the past century in the eastern U.S., and important causal factors often differ among them. Coincident with a recent decline event in upland oak-dominant forests of Arkansas, Missouri, and Oklahoma was an unexpected outbreak of a native cerambycid beetle, Enaphalodes rufulus (Haldeman), the red oak borer. A large range in estimates of oak mortality throughout affected forests was presumably due to variation in species composition, where oak-dominant areas experienced the greatest mortality. We chose eight sites across the Ozark and Ouachita National Forests of Arkansas, similar both topographically and by oak dominance, to determine if other stand or tree characteristics were important factors in variation of E. rufulus infestations across these forests. At each site, we sampled ∼125 dead, declining or healthy host Quercus rubra L., northern red oak. We created an estimate of the E. rufulus population level at each site during the recent outbreak using counts of dated larval gallery scars within a subset (n = 120) of all Q. rubra sampled (n = 976). We used classification tree partitioning to determine host tree characteristics that differed among dead, declining, and healthy Q. rubra. We also used classification tree partitioning, followed by logistic regression to determine stand characteristics that varied significantly among high, moderate and low infestation stands as well as between forests. Models indicated that trees which died were smallest, grew the least during the borer outbreak, and were apparently suppressed. These dying trees were likely poor competitors for resources, allowing neighboring survivors to experience a growth release during the E. rufulus outbreak. Larval survivorship was higher in trees which died, though larval densities were not greatest within these trees, which suggests that resistance in these individuals was compromised. At the stand level, differences between forests were apparently more important than those due to borer infestation. E. rufulus populations were higher at sites with lower Q. rubra basal area. This reduced basal area was likely a result of greater Q. rubra mortality at these sites during the borer outbreak in the early 2000s.     

Oak decline in the Boston Mountains,Arkansas, USA: Spatial and temporal patterns under two fire regimes

A spatially explicit forest succession and disturbance model is used to delineate the extent and dispersion of oak decline under two fire regimes over a 150-year period. The objectives of this study are to delineate potential current and future oak decline areas using species composition and age structure data in combination with ecological land types, and to investigate how relatively frequent simulated fires and fire suppression affect the dynamics of oak decline. We parameterized LANDIS, a spatially explicit forest succession and disturbance model, for areas in the Boston Mountains of Arkansas, USA. Land type distribution and initial species/age class were parameterized into LANDIS using existing forest data. Tree species were parameterized as five functional groups including white oak (Quercus alba L., Quercus stellata Wangenh., Quercus muehlenbergii Engelm.), red oak (Qurecus rubra L., Quercus marilandica Muenchh., Quercus falcata Michx., Quercus coccinea Muenchh.), black oak (Quercus velutina Lam.), shortleaf pine (Pinus echinata Mill), and maple (Acer rubrum L., Acer saccharum Marsh.) groups. Two fire regimes were also parameterized: current fire regime with a fire return interval of 300 years and a historic fire regime with an overall average fire return interval of 50 years. The 150-year simulation suggests that white oak and shortleaf pine abundance would increase under the historic fire regime and that the red oak group abundance increases under the current fire regime. The black oak group also shows a strong increasing trend under the current fire regime, and only the maple group remains relatively unchanged under both scenarios. At present, 45% of the sites in the study area are classified as potential oak decline sites (sites where red and black oak are >70 years old). After 150 simulation years, 30% of the sites are classified as potential oak decline sites under the current fire regime whereas 20% of the sites are potential oak decline sites under the historic fire regime. This analysis delineates potential oak decline sites and establishes risk ratings for these areas. This is a further step toward precision management and planning.     

Tree-cavity occurrence, cavity occupation and reproductive performance of secondary cavity-nesting birds in oak forests: The role of traditional management practices

Secondary cavity-nesting birds (SCN), which cannot create their own breeding cavities, are expected to be influenced by habitat alteration caused by forest management practices, but the mechanisms underlying the distribution pattern of SCN subjected to different management systems are poorly known. To improve our knowledge on these mechanisms, we examine cavity abundance, cavity occupation and reproductive performance of SCN in Pyrenean oak (Quercus pyrenaica) forests subjected to two management systems: (i) dense “young forests”, maintained at such stage by clear-cuttings and burns, and (ii) “old forest”, subjected to extensive traditional grazing and scarce firewood extraction by selective cutting. Young forests had considerably lower density of cavities (1.29 ± 0.71 vs 15.09 ± 2.00 cavities ha⁻¹), SCN species (0.18 ± 0.11 vs 0.61 ± 0.07 species ha⁻¹) and nests (0.40 ± 0.27 vs 2.67 ± 0.25 nests of all SCN ha⁻¹) than old forests, indicating that a low availability of cavities may limit SCN assemblages in young oak forests. However, reproductive parameters of great (Parus major) and blue (Cyanistes caeruleus) tits associated with the availability of food (laying date, clutch size, nestling number and weight, adult weight) did not differ between both forest types, suggesting that food supply was not reduced in young forests, at least for tits during the breeding season. Large diameter (up to 170 cm dbh) decayed trees were the most likely to hold cavities, but birds preferred smaller living cavity-trees for nesting (90% of nests in 21-65 cm dbh trees). The preservation of cavity-trees within traditionally managed old oak forests is crucial in providing nesting opportunities to SCN. Besides, the protection of these traditionally managed forests would also benefit to other forest organisms that depend on old and open oak forests.     

Monitoring the effects of gypsy moth defoliation on forest stand dynamics on Cape Cod,Massachusetts: Sampling intervals and appropriate interpretations

Gypsy moth (Lymantria dispar L.) outbreaks affect stand dynamics and successional patterns in pine–oak woodlands of North America's Atlantic coastal plain. Species-poor overstories are dominated by pitch pine (Pinus rigida), black oak (Quercus velutina) and white oak (Quercus alba). Both oaks are preferred food for gypsy moth, and white oak is the dominant late-successional species. We documented, over a 21-year period, the effects of gypsy moth defoliation on forest stand dynamics at Cape Cod National Seashore, Massachusetts. Our goal was to better understand the influence of gypsy moth defoliation on temporal changes in stand composition. Tree species abundance and frequency were measured in 16 stands with varying abundances of pitch pine and oak. Regression of 2-dimesional rates of change calculated from principal components analysis showed that gypsy moth significantly influenced stand dynamics and successional patterns between 1981 and 1992/1993. Defoliation was virtually absent in the 1990s, and analysis of 1992/1993–2002 rates of change showed topographic slope to be the most important factor influencing stand dynamics during this time. For the period 1981–2002 rates of change analysis failed to identify either factor as influencing changes in stand dynamics. We conclude that the 1980s defoliation events had a transitory effect on upland forest composition, but that there exists the potential for significant long-term impact due to recurrent defoliation episodes. Stand composition should be monitored at intervals which roughly correspond to the time it takes for stands to recover to disturbance events, in the case of those we sampled, approximately 10 years.     

Fine root dynamics of oak saplings in response to Phytophthora cinnamomi infection under different temperatures and durations

The belowground effects of Phytophthora cinnamomi on 1‐year‐old saplings of two common oak species in mid‐Atlantic US forests, white (Quercus alba) and red oak (Q. rubra), were examined after incubation in pathogen‐infested soilless potting mix. Fine root lengths (0–1.5 mm in diameter) of both oak species were quantified after incubation at successive 30‐day intervals up to 300 days, for a total of 10 incubation periods. In addition, colony‐forming units (CFU) of P. cinnamomi were quantified after white oak saplings were incubated in infested soilless potting mix at different temperature/duration combinations that reflect soil conditions present in the mid‐Atlantic United States. Impact of P. cinnamomi on fine root lengths of red and white oak saplings varied considerably over time. Significant periods of fine root loss occurred primarily during spring, when bud break and leaf flush began for both oak species. Red oaks had 17% fine root loss on average, while white oaks appeared more resistant to P. cinnamomi infection with a 2% decrease in fine roots over the course of the experiment. Phytophthora cinnamomi CFU declined significantly with exposure to all incubation temperatures except 8°C. This was in contrast to in vitro experiments, where the optimum temperature for mycelial growth was determined to be 21°C and above. Significant fine root loss caused by P. cinnamomi depended on plant phenology and the oak species tested. Extreme soil temperatures have a significant adverse impact on temporal changes of P. cinnamomi population.     

Water use by Tabor and Kermes oaks growing in their respective habitats in the Lower Galilee region of Israel

We estimated water use by the two main oak species of the Lower Galilee region of Israel—Tabor (Quercus ithaburensis) and Kermes (Quercus calliprinos)—to develop management options for climate-change scenarios. The trees were studied in their typical phytosociological associations on different bedrock formations at two sites with the same climatic conditions. Using the heat-pulse method, sap flow velocity was measured in eight trunks (trees) of each species during a number of periods in 2001, 2002 and 2003. Hourly sap flux was integrated to daily transpiration per tree and up-scaled to transpiration at the forest canopy level. The annual courses of daytime transpiration rate were estimated using fitted functions, and annual totals were calculated. Sap flow velocity was higher in Tabor than in Kermes oak, and it was highest in the youngest xylem, declining with depth into the older xylem. Average daytime transpiration rate was 67.9 ± 4.9 l tree⁻¹ d⁻¹, or 0.95 ± 0.07 mm d⁻¹, for Tabor oak, and 22.0 ± 1.7 l tree⁻¹d⁻¹, or 0.73 ± 0.05 mm d⁻¹, for Kermes oak. Differences between the two oak species in their forest canopy transpiration rates occurred mainly between the end of April and the beginning of October. Annual daytime transpiration was estimated to be 244 mm year⁻¹ for Tabor oak and 213 mm year⁻¹ for Kermes oak. Adding nocturnal water fluxes, estimated to be 20% of the daytime transpiration, resulted in total annual transpiration of 293 and 256 mm year⁻¹ by Tabor and Kermes oaks, respectively. These amounts constituted 51% and 44%, respectively, of the 578 mm year⁻¹ average annual rainfall in the region. The two species differed in their root morphology. Tabor oak roots did not penetrate the bedrock but were concentrated along the soil–rock interface within soil pockets. In contrast, the root system of Kermes oak grew deeper via fissures and crevices in the bedrock system and achieved direct contact with the deeper bedrock layers. Despite differences between the two sites in soil–bedrock lithological properties, and differences in the woody structure, annual water use by the two forest types was fairly similar. Because stocking density of the Tabor oak forests is strongly related to bedrock characteristics, thinning as a management tool will not change partitioning of the rainfall between different soil pockets, and hence soil water availability to the trees. In contrast, thinning of Kermes oak forests is expected to raise water availability to the remaining trees.     

Fire-driven oak regeneration in French Mediterranean ecosystems

Regeneration by seeds for cork oak (Quercus suber) and companion oaks (holm oak Quercus ilex and downy oak Quercus pubescens) is likely to be poor in the fire-prone Maures massif (southern France) but the causes are poorly known. Our objective was to assess the effective recruitment for these three oak species and their temporal pattern of recruitment, in order to determine the main limitation factors and the regeneration window of each species. We studied oak recruits (height <3 m) in naturally regenerated populations according to a gradient of fire recurrence and in five main vegetation types including shrublands and mixed mature woodlands. Fire recurrence was the main explanatory factor of oak recruitment, either directly or through vegetation type and microsite characteristics. The results indicate nil to low recruitment for holm oak and downy oak in shrublands, especially those recurrently burned and dominated by Cistus species. Cork oak recruited better than the other oaks in medium and high shrublands dominated by Erica arborea. In contrast, recruitment was high for holm and downy oak in mixed oak stands and mixed pine-oak stands that have not burned for decades. Microsite conditions such as coverage by litter and shrubs influenced oak recruitment, whereas landscape configuration and stand basal area had no influence. Our results suggest that strategic shrub-clearing, oak planting and protection of mixed oak woodlands as seed sources would help maintaining oak populations in the woodland–shrubland mosaic.     

Flying vs. climbing: Factors controlling arboreal seed removal in oak-beech forests

Nuts are heavy and nutritious seeds that need animals to be successfully dispersed. Most studies address nut removal by a single animal species once seeds fall onto the ground. However, nuts are also accessible before the seed drop and usually to a wide guild of seed foragers. This study examines the factors controlling arboreal seed removal in oak-beech forests within the whole guild of nut foragers. We found that seed-dispersing rodents (Apodemus sylvaticus) were the main acorn removers in the oaks (up to 3.75 m height), with a rapid seed encounter and a high removal rate. However, rodents did not climb the beech trees, probably due to their smoother bark in comparison to oak bark and/or the lower nutritional value of beechnuts with regard to acorns. Jays (Garrulus glandarius) were more abundant in oak stands (both dense and scattered) and clearly preferred acorns to beechnuts whereas nuthatches (Sitta europaea) were more abundant in beech stands and preferred beechnuts to acorns. Non-storing birds such as great tits (Parus major) also removed acorns and beechnuts, especially in the stands where oaks are dominant. Jays and rodents preferred sound seeds over insect-infested seeds but such a preference was not found for nuthatches. This study highlights that pure beech stands showed a reduced guild of arboreal nut foragers in comparison to oak stands. This different guild could probably affect the spatial patterns of seed dispersal, with a proportionally higher number of long dispersal events for acorns (mostly jay-dispersed) than for beechnuts (mostly nuthatch-dispersed). Long-distance dispersal of beechnuts (by jays) is determined by the presence of other preferred species (oaks) and their frequency of non-mast years. Seed location in different habitats strongly determines the contribution of different arboreal removers (including climbing rodents) and their removal speed, leading to a differential seed fate that will eventually affect tree regeneration. As nuthatches are sedentary birds, it is important to maintain old and dead trees where they can breed (crevices), forage (arthropods) and store seeds in order to favor beechnut dispersal and gene flow. By maintaining or favoring oak trees within beech stands we will ensure a wider guild of arboreal nut dispersers.     

Responses of oaks and tanoaks to the sudden oak death pathogen after 8 y of monitoring in two coastal California forests

Sudden oak death, caused by Phytophthora ramorum, is widely established in mesic forests of coastal central and northern California. In 2000, we placed 18 plots in two Marin County sites to monitor disease progression in coast live oaks (Quercus agrifolia), California black oaks (Q. kelloggii), and tanoaks (Lithocarpus densiflorus), the species that are most consistently killed by the pathogen in these areas. Through early 2008, the numbers of newly infected trees increased for all species. The infection rate for trees that were asymptomatic in 2000 was 5.0% y⁻¹ for coast live oaks, 4.1% y⁻¹ for black oaks and 10.0% y⁻¹ for tanoaks. Mortality rates were 3.1% y⁻¹ for coast live oaks, 2.4% y⁻¹ for black oaks, and 5.4% y⁻¹ for tanoaks. Mortality not attributed to P. ramorum was 0.54% y⁻¹ for coast live oaks, and 0.75% y⁻¹ for tanoaks. Weibull survival models of trees that were asymptomatic in 2000 provided overall median survival times of 13.7 y for coast live oaks, 13.8 y for black oaks, and 8.8 y for tanoaks. Survival of infected (bleeding) trees declined to 9.7 y for coast live oaks, 6.2 y for black oaks, and 5.8 y for tanoaks. Ambrosia beetle attacks on bleeding trees further reduced modeled survival times by 65–80%, reaffirming the earlier finding that beetle attacks on bleeding cankers considerably reduce survival. Across all plots, the modeled time for 90% of trees that were asymptomatic in 2000 to become infected is 36.5 y for coast live oaks and 15.4 y for tanoaks. There was a trend toward higher infection rates as tree diameter increased. Greater than 90% of living coast live oaks that failed during the study had extensive beetle tunneling at the site of the break. Disease intensity in coast live oaks at the plot level was positively associated with bay laurel (Umbellularia californica) basal area and negatively associated with Pacific madrone (Arbutus menziesii) basal area. This study demonstrates the use of survival modeling to characterize the effects of epidemic disease on different species and to project the future of forests infected with tree pathogens.     

Susceptibility of the Quercus rubra root system to Phytophthora cinnamomi; comparison with chestnut and other oak species

The susceptibility of oak seedlings (Quercus palustris, Quercus robur, Quercus rubra) and chestnut seedlings (Castanea sativa) to Phytophthora cinnamomi was tested. The dynamics of infection was examined in plant material raised in a rhizotron. In the oak species, primary root tissues were susceptible whereas secondary cortical tissues showed some resistance to P. cinnamomi. Secondary cortical tissues of the tap root in C. sativa were susceptible. Inoculations with P. cinnamomi were performed both in situ and on excised roots of mature Q. rubra. In both cases, the resistance of Q. rubra roots and shoots was negatively correlated with diameter at the inoculation point. Small roots (l–5-cm diameter) were resistant, whereas collar and trunk were susceptible. In contrast to oak, small excised roots of mature C. sativa (0.7–2-cm diameter) were susceptible to P. cinnamomi. This may explain why P. cinnamomi does not induce a decline of the attacked oaks, but rather a trunk canker.     

Oak (Quercus robur L.) regeneration in early successional woodlands grazed by wild ungulates in the absence of livestock

Wooded pastures grazed by livestock are believed to be landscapes that provide favourable conditions for spontaneous regeneration of oaks, including Quercus robur. A key mechanism for oak regeneration in these systems is ‘associational resistance’, spatial association with unpalatable plants which offer protection against herbivory. There is little knowledge on how oak regenerates without livestock grazing and in the presence of only wild large herbivores. We studied this in an area (114 ha) abandoned from agricultural use and in the early 1980s incorporated into the Bia?owie?a National Park, Poland. Its ungulate community consists of native red deer, European bison, roe deer, moose and wild boar. Secondary succession has led to the development of a mosaic habitat including tree and tall shrub groves (29% of the area), open meadow communities (60%), and edge, transitory zone between groves and meadows (11%). Our systematic inventory assigned oaks to height classes (0-0.2, 0.2-0.5, 0.5-1.3, 1.3-2.5, 2.5-5.0, >5.0 m), dichotomous shape characteristic (regular vs. “bonsai” sapling), as well as a habitat definition, in particular the characteristics of woody vegetation in the immediate surroundings of oaks. A selection of 17 oaks was subject to coring for the comparison of growth dynamics. Oak density was highest inside groves, with 504 oaks ha⁻¹, and in the edge zone (493 oaks ha⁻¹) and lowest in meadows (47 oaks ha⁻¹). Most of the 0-5-m oaks (62%) grew without another woody plant species within 1 m radius. The remaining oaks (38%) were associated mainly with Rubus idaeus and saplings of Carpinus betulus and Populus tremula - all highly ungulate-preferred species. The age (0.5 m above ground) of cored oaks in grove and edge habitats varied from 11 to 37 years, indicating continuous recruitment since agricultural abandonment. The initial growth dynamics of the more mature oaks did not differ from that of present “bonsais,” supporting the idea that browsing is not an unconditional impediment and that “bonsai” can be a temporary stage of successful oak development. In contrast to other studies, we found that associational resistance from unpalatable plants is not necessary to secure successful oak regeneration in woodlands subject to browsing by wild ungulates. This might have been possible because of the abundance of highly attractive vegetation making oak relatively unpreferred by ungulates. We suggest that the observed secondary succession provides a contemporary analogy of historic processes that resulted in the establishment of broadleaf forests with a substantial proportion of oak.     

Occurrence of Diplodia corticola,including new oak host records,in Wisconsin,USA

In the United States, diseased oaks (Quercus species) exhibit tip blight, branch and stem cankers, and dieback often attributed to Diplodia species or related fungi. Emergence of Diplodia corticola as a pathogen of European oaks, and reports of this fungus in the eastern and western United States, prompted re‐examination of strains from Wisconsin. These had been obtained in the late 1990s and early 2000s and previously identified only as Diplodia species. Nuclear ribosomal ITS sequences were obtained from the Wisconsin strains and analysed with other sequences from GenBank. Wisconsin strains confirmed as D. corticola were from northern red oak (Q. rubra), black oak (Q. velutina), white oak (Q. alba) and bur oak (Q. macrocarpa). Other strains from oaks in Wisconsin were D. mutila and D. seriata. Wound inoculation of northern red, white and bur oak seedlings with D. corticola in a greenhouse resulted in shoot death and stem lesions, from which the pathogen was reisolated. We conclude that D. corticola has been present in the northcentral United States for at least two decades and report two previously unrecognized hosts of this pathogen: white oak and bur oak. The roles of D. corticola, related fungi and influences of other environmental factors in deterioration of oak health in North America merit additional investigation.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.