The aftermath of an invasion: Structure and composition of Central Appalachian hemlock forests following establishment of the hemlock woolly adelgid, <Emphasis Type="Italic">Adelges tsugae</Emphasis>

As the highly invasive hemlock woolly adelgid, Adelges tsugae, continues to expand its distribution in eastern North America, affected forests will incur drastic changes in composition and structure. While these changes have been well-studied in dense hemlock forests in the Northeast, relatively little work is known about the effects of the adelgid at the western edge of the range of eastern hemlock, Tsuga canadensis. We evaluated the nature and extent of these changes using vegetation assessments coupled with growth simulations. The woody plant community was assessed in three strata (upper, mid- and lower) and was used to predict forest succession. Using the Forest Vegetation Simulator (FVS), we then projected the growth of hemlock forests 20 years into the future with and without the effects of the adelgid. In forest simulations lacking adelgid invasion, little change in composition or structure is forecast. In contrast, our projections predict a near complete loss of the hemlock forest type within 20 years of adelgid establishment, with widespread conversion to hardwood forest types, most notably white oak-red oak-hickory, chestnut oak-black oak-scarlet oak, and yellow poplar-white oak-red oak. Hemlock loss will result in denser deciduous forests with thinner canopies and multiple gaps, and significant alterations to terrestrial and aquatic wildlife habitat.     

Stand development on reforested bottomlands in the Mississippi Alluvial Valley

Reforestation of bottomland hardwood sites in the southeastern United States has markedly increased in recent years due, in part, to financial incentives provided by conservation programs. Currently >250,000 ha of marginal farmland have been returned to hardwood forests. I observed establishment of trees and shrubs on 205 reforested bottomlands: 133 sites were planted primarily with oak species (Quercus spp.), 60 sites were planted with pulpwood producing species (Populus deltoides, Liquidambar styraciflua, or Platanus occidentalis), and 12 sites were not planted (i.e., passive regeneration). Although oak sites were planted with more species, sites planted with pulpwood species were more rapidly colonized by additional species. The density of naturally colonizing species exceeded that of planted species but density of invaders decreased rapidly with distance from forest edge. Trees were shorter in height on sites planted with oaks than on sites planted with pulpwood species but within a site, planted trees attained greater heights than did colonizing species. Thus, planted trees dominated the canopy of reforested sites as they matured. Planted species acted in concert with natural invasion to influence the current condition of woody vegetation on reforested sites. Cluster analysis of species importance values distinguished three woody vegetation conditions: (1) Populus deltoides stands (2) oak stands with little natural invasion by other tree species, and (3) stands dominated by planted or naturally invading species other than oaks. Increased diversity on reforested sites would likely result from (a) greater diversity of planted species, particularly when sites are far from existing forest edges and (b) thinning of planted trees as they attain closed canopies.     

Driving factors of a vegetation shift from Scots pine to pubescent oak in dry Alpine forests

An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner‐Alpine regions, the species composition in low elevation forests is changing: The sub‐boreal Scots pine (Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub‐Mediterranean pubescent oak (Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger‐scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory (NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed‐effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small‐diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services.     

Modeling the Effects of Fire on the Long-Term Dynamics and Restoration of Yellow Pine and Oak Forests in the Southern Appalachian Mountains

We used LANDIS, a model of forest disturbance and succession, to simulate successional dynamics of forests in the southern Appalachian Mountains. The simulated environments are based on the Great Smoky Mountains landscapes studied by Whittaker. We focused on the consequences of two contrasting disturbance regimes—fire exclusion versus frequent burning—for the Yellow pine (Pinus L., subgenus Diploxylon Koehne) and oak (Quercus L.) forests that occupy dry mountain slopes and ridgetops. These ecosystems are a conservation priority, and declines in their abundance have stimulated considerable interest in the use of fire for ecosystem restoration. Under fire exclusion, the abundance of Yellow pines is projected to decrease, even on the driest sites (ridgetops, south‐ and west‐facing slopes). Hardwoods and White pine (P. strobus L.) replace the Yellow pines. In contrast, frequent burning promotes high levels of Table Mountain pine (P. pungens Lamb.) and Pitch pine (P. rigida Mill.) on the driest sites and reduces the abundance of less fire‐tolerant species. Our simulations also imply that fire maintains open woodland conditions, rather than closed‐canopy forest. For oaks, fire exclusion is beneficial on the driest sites because it permits oaks to replace the pines. On moister sites (north‐ and east‐facing slopes), however, fire exclusion leads to a diverse mix of oaks and other species, whereas frequent burning favors Chestnut oak (Q. montana Willd.) and White oak (Q. alba L.) dominance. Our results suggest that reintroducing fire may help restore decadent pine and oak stands in the southern Appalachian Mountains.     

Ecological succession following forest decline in a xeric oak forest of south-central United States

Aims The loss of canopy trees associated with forest decline can greatly influence the species composition and structure of a forest and have major impacts on the ecosystem. We studied the changes in forest composition and structure 1 and 5 years following nearly total canopy mortality on several hundreds of hectares of xeric oak forests in south-central United States. Because the forests were within an ecotonal vegetation type composed of a mosaic of forest, savanna and grassland, we sought to learn whether forest decline areas would recover to forest or change to more open savanna and grassland conditions in the landscape pattern of vegetation. Because low intensity fire shaped the vegetation type, we sought to learn whether fire would keep the decline areas open.Methods The study was conducted in a xeric oak forest in east-central Oklahoma, USA. Randomly located vegetation and regeneration surveys were conducted in decline and non-decline stands 1 and 5 years following nearly total canopy mortality. Diameter at breast height (DBH), regeneration and sprout origin were recorded for all woody species.Important findings The major canopy species post oak (Quercus stellata Wangenh.), blackjack oak (Quercus marilandica Muenchh.) and black hickory (Carya texana Buckl.) suffered 85–92% mortality; however, minor canopy components experienced limited mortality. Mortality affected all size classes of canopy trees except those below 5cm breast height diameter. There was abundant regeneration of all species and fire seemed to maintain a high level of sprouting. Decline appeared to decrease the relative importance of stump sprouting and increase other types including root sprouts. Decline areas had abundant true seedlings, with stem origin from a root with the same diameter as the stem, which is very unusual for xeric oak forests. Regeneration height in decline areas was twice that of non-decline forests. Our findings suggest that forest decline may lead to: (i) reduced oak dominance and species change in the canopy, (ii) change in reproduction type to increase success of true seedlings and maintain genetic diversity of oaks.     

Development of secondary pine forests after pine wilt disease in western Japan

Abstract. The development of secondary Pinus densiflora (Japanese red pine) forests after pine wilt disease was studied through phytosociological analysis, estimation of forest structure before disease and size-structure, tree ring and stem analyses. Following the end of the disease, the growth of previously suppressed small oak trees was accelerated. This is quite different from the development of forests following fire, which starts with the establishment of pine seedlings. Pine wilt disease shifted the dominance of secondary forests from Pinus densiflora to Quercus serrata oak forest. In pine forests, disturbance by fire is important for forest maintenance. In contrast, disturbance by pine wilt disease leads to an acceleration of succession from pine forest to oak forest.     

Impacts of anthropogenic disturbances on forest succession in the mid-montane forests of Central Himalaya

We studied the influence of anthropogenic drivers on the distribution and regeneration of tree species in vegetation at different stages of succession from grasslands to oak forests in mid-montane Central Himalaya. We found fire, grazing, and lopping as the main factors hindering a progressive successional regime towards a late-successional oak community. Succession was studied in five vegetation formations (grasslands, pine, pine–oak, open oak, and dense oak), with similar site conditions, representing a theoretical successional sequence from early- to late-successional stages. A structured survey with uniform distribution of sampling plots in the five selected vegetation formations was conducted to gather information abut the vegetation communities. Early-successional grasslands and pine forests were found to harbour high densities of pine and oak seedling and sapling regeneration. However, recurring fires and chronic unsustainable levels of grazing in these vegetation formations obstructed progressive succession by eliminating regenerating seedling and saplings from the forest understorey. Similarly, in intermediate- and late-successional stages (including pine–oak, open oak, and dense oak), overexploitation of existing oaks trees via lopping and grazing of regenerating oak seedlings and saplings hampered oak regeneration and development. The possibility to convert pine forests into oak as well as the conservation of existing oak forests through controlled grazing and lopping are management options that can contribute to an enhanced functioning of forest ecosystems in the study area. We conclude that with strategic management that restricts the current anthropogenic disturbances, the extent of oak forest in the study area can be increased.     

Projecting the distribution of forests in New England in response to climate change

Aim To project the distribution of three major forest types in the northeastern USA in response to expected climate change. Location The New England region of the United States. Methods We modelled the potential distribution of boreal conifer, northern deciduous hardwood and mixed oak–hickory forests using the process‐based BIOME4 vegetation model parameterized for regional forests under historic and projected future climate conditions. Projections of future climate were derived from three general circulation models forced by three global warming scenarios that span the range of likely anthropogenic greenhouse gas emissions. Results Annual temperature in New England is projected to increase by 2.2–3.3 °C by 2041–70 and by 3.0–5.2 °C by 2071–99 with corresponding increases in precipitation of 4.7–9.5% and 6.4–11.4%, respectively. We project that regional warming will result in the loss of 71–100% of boreal conifer forest in New England by the late 21st century. The range of mixed oak–hickory forests will shift northward by 1.0–2.1 latitudinal degrees (c. 100–200 km) and will increase in area by 149–431% by the end of the 21st century. Northern deciduous hardwoods are expected to decrease in area by 26% and move upslope by 76 m on average. The upslope movement of the northern deciduous hardwoods and the increase in oak–hickory forests coincide with an approximate 556 m upslope retreat of the boreal conifer forest by 2071–99. In our simulations, rising atmospheric CO₂ concentrations reduce the losses of boreal conifer forest in New England from expected losses based on climatic change alone. Main conclusion Projected climate warming in the 21st century is likely to cause the extensive loss of boreal conifer forests, reduce the extent of northern hardwood deciduous forests, and result in large increases of mixed oak–hickory forest in New England.     

Pathogen induced disturbance and succession in temperate forests: Evidence from a 100-year data set in southern Sweden

Major tree species are declining in many temperate forests due to changing disturbance regimes, including invasive pests and pathogens. We examined the interaction of secondary succession and Dutch elm disease in the Swedish temperate forest reserve Dalby Söderskog, based on five tree surveys made between 1909 and 2011. The forest is characterized by the coexistence of four major European tree species: wych elm (Ulmus glabra), European ash (Fraxinus excelsior), European beech (Fagus sylvatica) and pedunculate oak (Quercus robur). After protection of the forest in 1918, lack of disturbance mainly favoured elm, while the oak population declined due to mortality of old oaks and lack of regeneration. Dutch elm disease has caused high and continuous elm mortality since 1988. As a result, increased light availability at the forest floor favoured abundant regeneration of ash, beech, and lately also oak. The recent arrival of an invasive fungal pathogen causing ash dieback may once again change the course of succession. Open space emerging from loss of elm and ash in forest reserves may be used by reserve managers to favour oak regeneration and biodiversity of semi-open woodlands once lost during succession to closed forest. We conclude that winners and losers change places as an effect of invasive pathogens, resulting in unexpected successions and both losses and gains in valuable ecological niches and habitat structures in temperate broadleaf forests.     

Long‐term directional changes in upland Quercus forests throughout Oklahoma,USA

Questions: (1) How have the composition and structure of undisturbed upland Quercus forests changed over 50 years across a large region and moisture gradient; (2) What factors are associated with long‐term and broad‐scale changes in these forests? Location: Oklahoma, USA. Methods: We re‐sampled 30 forest stands originally sampled in the 1950s across a large geographical area and compared basal area, tree density, and sapling density between the sampling periods using paired t‐tests, CCA, and DCA. We examined vegetation dynamics in the context of drought indices compiled for the sample period. Results: Total and Quercus stellata basal area and tree density increased, but Q. stellata and Q. marilandica sapling density decreased. Juniperus virginiana and woody species richness increased for all measures. DCA indicated that re‐sampled stands generally changed from Q. stellata–Q. marilandica‐dominated forests to forests with greater woody species richness and more J. virginiana. Q. stellata remained a dominant tree species; otherwise, composition shifted towards mesophytic and invasive woody species. Measurements taken in the 1950s immediately followed a major drought; whereas subsequent decades were significantly moister. Conclusions: Fire exclusion and drought may have played an important role in driving changes towards lower dominance by Quercus, increased importance of mesophytic and invasive species, and greater woody species richness. These phenomena are similar to those found in Quercus‐dominated forests throughout the northern hemisphere.     

Colonization, survival, and causes of mortality of Camerariu hamadryadella (Lepidoptera: Gracillariidae) on four species of host plants

Abstract.

• 1 The abundance, survival, and causes of mortality of Cameraria hamadryadella (Clemens) (Lepidoptera: Gracillariidae) were examined on four host plant species in Virginia, U.S.A. Quercus alba L. and Q.rubra L. are native within the geographic range of C.hamadryadella, and Q.robur L. and Q.benderi Baenitz are exotic. Q.robur is native to Europe, North Africa, and Asia and was probably introduced prior to 1850, and Q.benderi is of hybrid origin and introduced to cultivation before 1900. Q.alba and Q.robur are in the subgenus Lepidobalanus (white oaks), and Q.rubra and Q.benderi are in the subgenus Erythrobalanus (red oaks).
• 2 Larval mines of C. hamadryadella were abundant on both white oak species, including the exotic Q.robur, but were rare on host plants in the red oak subgenus. Un-hatched eggs of C.hamadryadella were not observed on red oaks. Other observations on host distribution indicate that C.hamadryadella is rarely found on red oaks. These observations are interpreted as circumstantial evidence that C. hamadryadella generally avoids ovipositing on red oaks.
• 3 Survival of C.hamadryadella to the adult stage was similar among all host species, but larvae tended to survive longer on hosts in the red oak subgenus. The observation of higher survival rates of early instar larvae on red oaks suggests that no nutritional or secondary chemical barrier reinforces the observed pattern of oviposition.
• 4 Significant differences in the distribution of the causes of mortality were detected between native and exotic host plant species. Larvae and pupae on native hosts were more likely to die because of predation, while those on exotic host plants were more likely to die because of parasitism and host feeding by adult female parasitoids. This pattern could arise because parasitoids prefer to forage on exotic host plants or because predators avoid foraging on exotic plants.
• 5 This study shows for C. hamadryadella that the only barriers to colonization and use of exotic hosts, in the white and red oak subgenera, are the presence of cues sufficient to stimulate oviposition and/or the absence of cues to deter oviposition. It also suggests that the presence of closely related native host plants in the region of introduction will increase the probability that exotic plants will be colonized by phytophagous insects.

     

Effects of competition and indirect facilitation by shrubs on <Emphasis Type="Italic">Quercus robur</Emphasis> saplings

Indirect facilitation by shrubs has been suggested as a cost-effective way of regenerating oaks in forests of conservation interest. In this study, we tested whether shrubs can enhance growth in pedunculate oak (Quercus robur) by suppressing herbaceous competitors. We studied interactions between young oaks, shrubs, and/or herbaceous vegetation in an open-field experiment, in southern Sweden, over the first 3 years after planting. Oak saplings were grown in four competition treatments: no competing vegetation; with herbaceous vegetation; with shrubs; and with both herbaceous vegetation and shrubs. Competition from shrubs and herbaceous vegetation both reduced stem diameter and biomass accumulation, but they affected biomass partitioning differently. Saplings grown with competition from shrubs partitioned biomass primarily into height growth, while those saplings exposed to competition from herbaceous vegetation invested a relatively higher proportion in root growth. Competition between shrubs and herbaceous vegetation reduced the above-ground biomass of the herbaceous vegetation, resulting in an indirect facilitative effect for the oaks during the first 2 years after planting. However, during the third year, shrubs had a negative effect on biomass accumulation. In summary, results from this study suggest that shrubs indirectly facilitate biomass accumulation of oak saplings by suppressing herbaceous vegetation, possibly by reducing competition for below-ground resources. However, owing to the relatively short duration of positive net outcome for the oak, we recommend that a longer-term assessment of the interaction between oak regeneration and neighboring shrubs be made before the outcome of this study is applied to practical forestry.     

Hollow oaks and beetle conservation: the significance of the surroundings

In this study we investigated hollow oaks (Quercus robur, Q. petrea) situated in open landscapes and in forests in Norway in northern Europe, and compared their importance for rare and threatened beetles (Coleoptera). Old, hollow oak trees, both in parks and in forests, were extremely rich in red-listed beetles, and hosted a high proportion of threatened species. The proportion of oak associated species and the mean number of red-listed beetle species per tree was similar in the two site types, but rarefaction showed that for a certain number of individuals, oaks in forests had more threatened and near-threatened species than oaks in parks. The species composition also differed between site types: Park oaks had a higher proportion of species associated with hollows and animal nests, whereas in forests, there was a higher proportion of species depending on dead oak wood in general. Four factors were significant in explaining the richness of red-listed beetles in our study: Tree circumference, cavity decay stage, proportion of oak in the surroundings, and coarse woody debris (CWD) in the surroundings. Forest oaks were smaller, but they still trapped a species richness comparable to that of the larger park oaks—probably a result of high amounts of CWD in the surroundings. We show that oaks in open landscapes and oaks in forest have only partly overlapping beetle assemblages and, thus, cannot be substituted in conservation. Planning for conservation of red-listed beetles associated with this key habitat demands a large scale perspective, both in space and time, as the surroundings have important effects on associated threatened and near threatened species.     

Springtime in the city: exotic shrubs promote earlier greenup in urban forests

Despite the widespread recognition that urban areas are frequently dominated by exotic and invasive plants, the consequences of these changes in community structure have not been explicitly considered as an explanation for the pattern of advanced leaf phenology, or early greenup, reported in many urban areas. As such, we evaluated two hypotheses that could account for advanced greenup in forests along an urban to rural gradient: advanced phenology within individual species or differences in woody plant community. We monitored the spring leafing phenology of Aesculus glabra (Ohio buckeye), Lonicera maackii (Amur honeysuckle), and Acer negundo (box elder) in 11 forests spanning an urban to rural gradient in central Ohio, USA. From February to April 2006, we monitored these species, recorded woody plant composition, and documented daily minimum and maximum temperatures at each site. We found a weak but general trend of advanced phenology within species in more urban landscapes. Monthly average minimum temperatures were higher with increasing urbanization while monthly average maximum temperatures were similar across the urban to rural gradient. We also found evidence for shifts in woody plant communities along the urbanization gradient, mainly driven by the abundance of L. maackii, an invasive exotic species, in the more urban forests. Because L. maackii leafs out weeks earlier than native woody species and is very abundant in urban forests, we suggest that the invasion of forests by this species can generate earlier greenup of urban forests.     

Influence of canopy openness,ungulate exclosure,and low‐intensity fire for improved oak regeneration in temperate Europe

Failed oak regeneration is widely reported in temperate forests and has been linked in part to changed disturbance regimes and land‐use. We investigated if the North American fire–oak hypothesis could be applicable to temperate European oaks (Quercus robur, Quercus petraea) using a replicated field experiment with contrasting canopy openness, protection against ungulate browsing (fencing/no fencing), and low‐intensity surface fire (burn/no burn). Survival, relative height growth (RGR_H), browsing damage on naturally regenerated oaks (≤300 cm tall), and changes in competing woody vegetation were monitored over three years. Greater light availability in canopy gaps increased oak RGR_H (p = .034) and tended to increase survival (p = .092). There was also a trend that protection from browsing positively affected RGR_H (p = .058) and survival (p = .059). Burning reduced survival (p < .001), nonetheless, survival rates were relatively high across treatment combinations at the end of the experiment (54%–92%). Most oaks receiving fire were top‐killed and survived by producing new sprouts; therefore, RGR_H in burned plots became strongly negative the first year. Thereafter, RGR_H was greater in burned plots (p = .002). Burning altered the patterns of ungulate browsing frequency on oaks. Overall, browsing frequency was greater during winter; however, in recently burned plots summer browsing was prominent. Burning did not change relative density of oaks, but it had a clear effect on competing woody vegetation as it reduced the number of individuals (p < .001) and their heights (p < .001). Our results suggest that young, temperate European oaks may respond similarly to fire as their North American congeners. However, disturbance from a single low‐intensity fire may not be sufficient to ensure a persistent competitive advantage—multiple fires and canopy thinning to increase light availability may be needed. Further research investigating long‐term fire effects on oaks of various ages, species‐specific response of competitors and implications for biodiversity conservation is needed.     

Multiple Recruitment Limitation Causes Arrested Succession in Mediterranean Cork Oak Systems

Lack of tree regeneration and persistency of species-poor shrublands represent a growing problem across Mediterranean evergreen oak forests. What constrains forest regeneration is poorly understood, and restoration attempts have been largely unsuccessful. We assessed the contribution of four different mechanisms of tree recruitment limitation (that is, source, dispersal, germination, and establishment) in a cork oak (Quercus suber) system in southern Portugal. Using a combination of field studies and experiments, we quantified seed production, seed removal and dispersal, seed survival and germination, seedling establishment and survival, as well as cork oak natural regeneration for the three dominant vegetation types in this system (Cistus ladanifer shrubland, oak forest, and oak savanna). We found that all four forms of cork oak recruitment limitation were significantly more severe in shrublands than in oak forests and savannas, so that oak seedling recruitment in shrubland was impeded in multiple ways. Our results explain why transitions from shrublands to oak savannas and forests are extremely difficult, and that the release from arrested succession in this system requires the simultaneous relief of multiple constraints on recruitment limitation in the early life history of oaks. These results have important implications for the restoration and conservation of Mediterranean oak systems.     

Carbon density and distribution of six Chinese temperate forests

Quantifying forest carbon (C) storage and distribution is important for forest C cycling studies and terrestrial ecosystem mod-eling. Forest inventory and allometric approaches were used to measure C density and allocation in six representative temper-ate forests of similar stand age (42–59 years old) and growing under the same climate in northeastern China. The forests were an aspen-birch forest, a hardwood forest, a Korean pine plantation, a Dahurian larch plantation, a mixed deciduous forest, and a Mongolian oak forest. There were no significant differences in the C densities of ecosystem components (except for detritus) although the six forests had varying vegetation compositions and site conditions. However, the differences were significant when the C pools were normalized against stand basal area. The total ecosystem C density varied from 186.9 tC hm^-2 to 349.2 tC hm^-2 across the forests. The C densities of vegetation, detritus, and soil ranged from 86.3–122.7 tC hm^-2, 6.5–10.5 tC hm^-2, and 93.7–220.1 tC hm^-2, respectively, which accounted for 39.7%±7.1% (mean±SD), 3.3%±1.1%, and 57.0%±7.9% of the total C densities, respectively. The overstory C pool accounted for > 99% of the total vegetation C pool. The foliage bio-mass, small root (diameter < 5mm) biomass, root-shoot ratio, and small root to foliage biomass ratio varied from 2.08–4.72 tC hm^-2, 0.95–3.24 tC hm^-2, 22.0%–28.3%, and 34.5%–122.2%, respectively. The Korean pine plantation had the lowest foliage production efficiency (total biomass/foliage biomass: 22.6 g g^-1) among the six forests, while the Dahurian larch plantation had the highest small root production efficiency (total biomass/small root biomass: 124.7 g g^-1). The small root C density de-creased with soil depth for all forests except for the Mongolian oak forest, in which the small roots tended to be vertically dis-tributed downwards. The C density of coarse woody debris was significantly less in the two plantations than in the four natu-rally regenerated forests. The variability of C allocation patterns in a specific forest is jointly influenced by vegetation type, management history, and local water and nutrient availability. The study provides important data for developing and validating C cycling models for temperate forests.     

Modelling Mediterranean oak palaeolandscapes using the MaxEnt model algorithm: The case of the NE Iberia under the Middle Holocene climatic scenario

The Mediterranean Basin is a global biodiversity hotspot, and oak tree species play an important role in it. Since the beginning of the Holocene (∼11.4 kyr BP), the distribution of forests has not occurred uniformly, resulting in diverse types of vegetation landscapes. In this study, we used a maximum entropy algorithm (MaxEnt) to obtain the ecological niche model (ENM) of two sub-Mediterranean oak species, Quercus pubescens Willd. (pubescent oak) and Quercus ilex subsp. ilex (holm oak), both in the present day in the Iberian Peninsula and within a Middle Holocene (8.2–4.2 kyr BP) climatic scenario in the NE Iberian Peninsula. Moreover, we used the locations of Neolithic archaeological sites containing anthracological data to analyze the relationship between human occupations and oak habitats. Our results suggest that the two oaks have responded differently to the climatic conditions that have occurred, and show changes in both potential distributions. The palaeolandscape vegetation map shows a denser vegetation cover in the lowlands and a more open landscape in the highlands, with a higher dominance of Quercus pubescens in the septentrional areas, while Quercus ilex was more restricted to certain coastal areas. Temperature and precipitation factors, mainly seasonal climatic conditions, have had a greater impact on the distribution of vegetation than other factors. We found a good overlap between the ENM of the two oaks and the locations of the Neolithic sites analysed, and determined that the distribution of Neolithic archaeological sites is not random. The Neolithic populations in the study area depended heavily on the resources of the deciduous and evergreen sub- Mediterranean forest, although they also exploited the resources of the mountain pine forest. Neolithic sites distribution suggests that Neolithic human groups were aware of the potential of forests and probably gathered woody resources in their surroundings.     

Densification and State Transition Across the Missouri Ozarks Landscape

World-wide, some biomes are densifying, or increasing in dense woody vegetation, and shifting to alternative stable states. We quantified densification and state transition between forests ecosystems in historical (ca. 1815–1850) and current (2004–2008) surveys of the Missouri Ozark Highlands, a 5-million ha landscape in southern Missouri, USA. To estimate density of historical forests, we used the Morisita plotless density estimator and applied corrections for surveyor bias. For contemporary forests, we used known densities at plots to predict continuous densities with random forests, an ensemble regression tree method. We also calculated basal area and percent stocking to determine changes in wood volume. Historical forests had densities ranging from about 75 to 320 trees/ha. Current forest densities were about 2.3 times greater and more uniform, at about 300–400 trees/ha (DBH ≥ 12.7 cm). Not all forests have increased in basal area and percent stocking because trees in contemporary forests are smaller in diameter than historical forests. Although oak species still are dominant (as defined by ≥10% composition), oak dominance is being replaced by many fire-sensitive species, of which only eastern redcedar and maples have become dominant. Densification and community changes in functional traits have produced a state transition from open oak forest ecosystems to predominantly closed eastern broadleaf forests in the Missouri Ozarks. This shift is not at equilibrium, as fire-sensitive species are continuing to increase and turnover in long-lived oaks is slow.     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.