Modeling the effects of alternative management strategies on forest carbon in the Nothofagus forests of Tierra del Fuego,Chile

The impact of forest management activities on the ability of forest ecosystems to sequester and store atmospheric carbon is of increasing scientific and social concern. The nature of these impacts varies among forest ecosystems, and spatially and temporally explicit ecosystem models are useful for quantifying the impacts of a number of alternative management regimes for the same forest landscape. The LANDIS-II forest dynamics simulation model is used to quantify changes to the live overstory and coarse woody debris pools under several forest management scenarios in a high-latitude South American forest landscape dominated by two species of southern beech, Nothofagus betuloides and N. pumilio. Both harvest type (clearcutting vs. partial overstory retention) and rotation length (100 years vs. 200 years) were significant predictors of carbon storage in the simulation models. The prompt regeneration of harvest units greatly enhanced carbon storage in clearcutting scenarios. The woody debris pool was particularly sensitive to both harvest type and rotation length, with large decreases noted under short rotation clearcutting. The roles of extended rotations and partial overstory retention are noted for enhancing net carbon storage on the forest landscape.     

Forest management conditioning ground ant community structure and composition in temperate conifer forests in the Pyrenees Mountains

The search for indicators to monitor management impact on biodiversity is a crucial question because management practices promote changes in community structure and composition of different animal groups. This study explores the effect of widely conducted management practices (forest logging and livestock) in Pinus uncinata forests in the Pyrenees range (NE Spain) on the structure and composition of ground ant communities compared to those of old-growth stands. Forest structure clearly differed in stands with different forest managements. These stands managed for different uses also showed marked differences in structure and composition of ground ant communities. There was a great dominance of a single species, Formica lugubris, which accounted for 99% of ants collected in pitfall traps. Rarefaction curves indicated that species richness was highest in old-growth stands and lowest in even-aged ones, with woodland pasture stands showing an intermediate value. Classification methods allowed us to identify two groups of species: six species related to old-growth plots and three species (including F. lugubris) associated to managed stands. Habitat structure played an important role in determining the structure of ant communities: forests with high tree density but low basal area were the most favourable forest type for F. lugubris, while the abundance of the remaining ant species was negatively affected by the abundance of F. lugubris and by tree cover.     

Climate and historical stand dynamics in the tropical pine forests of northern Thailand

Forest recruitment is the outcome of local- and regional-scale factors such as disturbances and climate. The relative importance of local- and regional-scale factors will determine the spatial scale at which temporal pulses of recruitment occur. In seasonal tropical forests, where the annual dry-season is a critical bottleneck to seedling survival, multi-year periods of relatively cool, wet dry seasons may be required for successful tree recruitment. Consequently, when such conditions are present, region-wide synchronisation of recruitment may occur. To examine the case for regional synchronisation of forest dynamics in the seasonal tropical pine forests of northern Thailand, we investigated forest age structures at three spatial scales: stand, site and region. We compared forest age structures with instrumental climatic records beginning in 1902. We found significant statistical evidence of synchronous recruitment at the stand- and site-scales, but not at the regional-scale. While correlations between recruitment and climate were not statistically significant, recruitment success was often linked to favourable climatic conditions. For example, recruitment at all sites was associated with multi-year periods of cool-wet dry seasons. The lack of significant correlations between recruitment and climate appears to reflect complex interactions among local disturbance history, regional climate variability and pine recruitment.     

Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.     

The lateral spread of tree root systems in boreal forests: Estimates based on N uptake and distribution of sporocarps of ectomycorrhizal fungi

In nutrient poor environments, such as boreal forests, many of the most important interactions between plants take place belowground. Here, we report the results of two approaches to obtain estimates of the lateral spread of tree roots.     

Restoration of dry tropical forests in Central America: A review of pattern and process

Much information on restoration and management exists for wet tropical forests of Central America but comparatively little work has been done in the dry forests of this region. Such information is critical for reforestation efforts that are now occurring throughout Central America. This paper describes processes of degradation due to land use and provides a conceptual framework for the restoration of dry tropical forest. Most of this forest type was initially harvested for timber and then cleared for cattle in the last century (1930-1970). Only 1.7% remains largely restricted to infertile soils and remote areas on the Pacific coastal side of Panama, Costa Rica, Nicaragua and Mexico. These cleared areas are again in a state of transition due to a combination of decreasing land productivity, and land speculation for tourism development. Some farms have been sold to new landowners who are interested in reforesting to increase biodiversity and forest cover. Attempts have therefore been made to reforest by protecting the land from fire and cattle, by supplementing natural regrowth with enrichment planting, or through use of tree plantations. Experimental studies have demonstrated the ability of these lands to grow back to forests because of native species ability to sprout after cutting, and the capacity of remnant trees in field and riparian zones to provide seeds and to moderate edge environment for seed germination and seedling establishment. However, research also shows that on sites with long histories of land clearance, species diversity will remain low with functional groups missing unless some active management occurs. Under-planting with late-successional native tree species can add structure and diversity; enrichment planting with large-fruited shade-intolerant species can initiate new islands of more diverse regeneration beneath their canopies; and plantings of fast-growing, nitrogen-fixing trees that provide light canopy shade can moderate the environment below, promoting regeneration establishment of late-successional species. Plantations are the only option for lands that have lost almost all remnants of native forest, and where soils and vegetation have changed to new states of structure and function. Conversion of pastures to tree plantations that can facilitate natural regeneration beneath them is appropriate when pastures are prone to fire and/or lack immediate seed sources nearby. After the grasses have been shaded out, natural recruitment can slowly occur over a 10-15 years period. Under-planting of shade-tolerant late-successional species can supplement species composition and structure.     

Driving factors for natural tree rejuvenation in anthropogenic pine (Pinus sylvestris L.) forests of NE Germany

The rejuvenation ecology of three main tree species in anthropogenic pine (Pinus sylvestris L.) forests is explored in our study. We focus on the scale of micro-plots, which provide the safe sites for tree rejuvenation. We thrive on the multi-factorial relationship of tree establishment and driving ecological factors using a large dataset from pine stands in NE Germany and applying multivariate analyses. The success of the establishment of the investigated focal tree species Fagus sylvatica L., Quercus petraea Liebl. and Pinus sylvestris L. is, on general, mostly affected by three factors, i.e. water balance of the upper soil layers, browsing pressure, and diaspore sources. Our investigations on the micro-plot scale revealed species-specific differences. For beech saplings <50 cm growth height, primarily the availability of water, indicated by available water capacity (AWC), thickness, quality, and structure of the organic layer, silt and humus content in the topsoil, and the lack of a dense competitive herb layer, were identified as most important factors. On the contrary, oak seems hardly be restricted by hydrologic and/or trophic deficits in the topsoil or humus layer. In conclusion and comparison to Fagus sylvatica L., we assume for Quercus petraea Liebl. advantages in natural regeneration processes under sub-continental climate conditions and thus under the scenarios of climate change. Pinus sylvestris L. regeneration in our investigation area occurs only in a narrow niche. We conclude with regard to future forest development and the objective of stand conversion with low management intensity that oak should be favoured within natural stand regeneration.     

Understory succession in post-agricultural oak forests: Habitat fragmentation affects forest specialists and generalists differently

The herbaceous understory forms the richest stratum in temperate broadleaved forests in terms of plant diversity. Understanding the process of understory succession is thus of critical importance for the development of management guidelines for biodiversity restoration in post-agricultural plantation forests.We studied effects of stand age, forest fragmentation, and soil and canopy conditions on species richness and abundance of four species groups in the understory of post-arable oak plantations in southern Sweden: herbaceous forest specialists, habitat generalists and open-land species, and woody species.The group of forest specialists may approach the richness of continuously forested sites after 60-80 years in non-fragmented plantations, but many forest species were sensitive to habitat fragmentation. Open-land species richness decreased during succession, while the richness of woody species and of generalists remained stable, and was not affected by fragmentation. Abundance of generalists gradually decreased in non-fragmented plantations, probably due to competition from colonizing forest specialists. Soil pH in post-arable stands remained consistently higher than in continuously forested stands, which maintained differences in species composition. The development of a shrub layer seemed to imply a competitive advantage for forest specialists compared to generalist species.For successful recovery of a rich understory, we suggest that post-arable plantations should be established on loamy soils of intermediate to high pH proximate to older forest with source populations, and that a continuous overstory canopy cover of 70-80% is maintained by regular light thinnings and promotion of a shrub layer.     

Diversity and structure of regenerating tropical dry forests in Costa Rica: Geographic patterns and environmental drivers

Much of the dry tropical forest biome has been converted to agricultural land uses over the past several centuries. However, in conservation areas such as those in the Guanacaste and Tempisque regions of Costa Rica, tropical dry forests are regenerating due to management practices including fire suppression. To better understand the patterns of secondary succession occurring in Costa Rican tropical dry forest, we established 60 20 × 50 m plots in mature and regenerating forests in the Sector Santa Rosa (formerly known as Parque Nacional Santa Rosa) and Palo Verde National Park. Plots were stratified into three plant communities: tropical dry oak forest (Quercus oleoides) (SROAK), Santa Rosa tropical dry forest (SRTDF), and Palo Verde tropical dry forest (PVTDF). In these plots we measured and identified and all individuals >10 cm DBH, measured but did not identify stems <10 cm but taller than 1.3 m, counted woody seedlings (<1.3 m height) and analyzed soil chemical and physical properties.     

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.     

Forest stand dynamics and sudden oak death: Mortality in mixed-evergreen forests dominated by coast live oak

Sudden oak death (SOD), caused by the recently discovered non-native invasive pathogen, Phytophthora ramorum, has already killed tens of thousands of native coast live oak and tanoak trees in California. Little is known of potential short and long term impacts of this novel plant–pathogen interaction on forest structure and composition. Coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica) form mixed-evergreen forests along the northern California coast. This study measured tree mortality over a gradient of disease in three time periods. Direct measurements of current mortality were taken during 2004, representing a point-in-time estimate of present and ongoing mortality. Past stand conditions, c. 1994, were estimated using a stand reconstruction technique. Future stand conditions, c. 2014, were calculated by assuming that, given a lack of host resistance, live trees showing signs of the disease in 2004 would die. Results indicate that coast live oaks died at a rate of 4.4–5.5% year⁻¹ between 1994 and 2004 in highly impacted sites, compared with a background rate of 0.49% year⁻¹, a ten-fold increase in mortality. From 2004 to 2014, mortality rates in the same sites were 0.8–2.6% year⁻¹. Over the entire period, in highly impacted sites, a 59–70% loss of coast live oak basal area was predicted, and coast live oak decreased from 60% to 40% of total stand basal area, while bay laurel increased from 22% to 37%. Future stand structures will likely have greater proportions of bay laurel relative to coast live oak.     

A century of riparian forest expansion following extreme disturbance: Spatio-temporal change in Populus/Salix/Tamarix forests along the Upper San Pedro River,Arizona, USA

Populus–Salix forests are a valued riparian vegetation type in western North America. These pioneer, obligate phreatophytes have declined on some rivers, raising conservation concerns and stimulating restoration plantings, but have increased on others. Understanding patterns and causes of forest change is essential for formulating conservation, restoration and management plans. Our goal was to assess spatio-temporal patterns of vegetation change on the Upper San Pedro River in semiarid Arizona, USA, one of the few undammed rivers in the region. Over 100 years ago, intense floods initiated channel incision and substantially altered hydrogeomorphology. Pioneer trees began to establish in the widening post-entrenchment zone as the surfaces began to stabilize. Using a time-series of aerial photographs (1955–2003) we quantified recent change in area of riparian cover types. Analysis indicated that wooded area in the post-entrenchment zone nearly tripled from 1955 to 2003, at the expense of bare ground, and the active channel narrowed appreciably. This forest expansion represents a long-term response to river entrenchment, with the temporal pattern influenced by recent flood cycles and biogeomorphic feedbacks. Populus–Salix have established episodically during the infrequent years with high winter flood runoff, sequentially filling available recruitment space. Older cohorts cover wide swaths of the floodplain while young trees form narrow bands lining the channel. Barring extreme flooding, the pioneer forests are expected to senesce over the coming century. An additional factor that has shaped the pattern of post-entrenchment forest expansion is anthropogenic water withdrawal. Populus–Salix forest increase has been greatest within a conservation area, where stream flows are largely perennial. In drier, agricultural sectors, Populus–Salix have declined while the more deeply-rooted Tamarix has increased. Overall, the study reveals that long-term fluctuations in pioneer forest area and age structure are common on dryland rivers, and shows how past events such as extreme floods can interact with recent environmental practices such as freshwater withdrawal to influence riparian forest patterns. This underscores the necessity of a long-term perspective for forest conservation and management.     

Long-term browsing impact around diversionary feeding stations for moose in Southern Norway

Diversionary winter feeding of browsing ungulates is an increasingly common management practice although evidence for its efficacy to reduce excessive browsing remains ambiguous. Moreover, comparative estimates on changes in browsing pressure (proportion of available shoots browsed) following long-term winter feeding are currently lacking. We quantified spatiotemporal changes in browsing pressure of moose (Alces alces L.) on commercial and non-commercial tree species around 30 feeding stations after 15–20 years of winter feeding. The results were compared with browsing pressure indices recorded at the same feeding stations 10 years previously. We expected leader stem and lateral twig browsing to have increased over time at a fine spatial scale (≤200 m from feeding station) and to have increased in spatial extent away from feeding stations. Furthermore, we tested whether moose browsing patterns conformed to central-place foraging theory. Despite 2–3-fold higher faecal pellet group numbers in the vicinity of feeding stations, leader stem browsing increased only on the commercially valuable Norway spruce (Picea abies (L.) Karst), a species normally avoided by moose. Lateral twig browsing largely decreased within 200 m of feeding stations while at a broader scale (≤1 km from feeding stations), leader stem browsing was high on most tree species (ca. 60% of available stems browsed) as expected, Peak browsing on lateral twigs of Scots pine (Pinus sylvestris L.) occurred further from feeding stations after 15–20 years of winter feeding than 10 years earlier. Browsing pressure on active feeding stations (n = 18) was comparable to that on inactive feeding stations (n = 12), suggesting the occurrence of rebrowsing. The ability of central-place foraging theory to explain fine-scale browsing patterns around feeding stations decreased as diversionary feeding continued over time. Long-term diversionary winter feeding of moose in unproductive boreal forests has serious implications for the intensity of fine-scale browsing pressure, which may lead to resource depletion close to feeding stations followed by high browsing pressure at distances further away from feeding stations. Our study emphasizes the importance of considering the longevity of a winter feeding programme at its inception.     

Epiphyte diversity and biomass loads of canopy emergent trees in Chilean temperate rain forests: A neglected functional component

We document for the first time the epiphytic composition and biomass of canopy emergent trees from temperate, old-growth coastal rainforests of Chile (42°30′S). Through tree-climbing techniques, we accessed the crown of two large (c. 1 m trunk diameter, 25–30 m tall) individuals of Eucryphia cordifolia (Cunoniaceae) and one large Aextoxicon punctatum (Aextoxicaceae) to sample all epiphytes from the base to the treetop. Epiphytes, with the exception of the hemi-epiphytic tree Raukaua laetevirens (Araliaceae), were removed, weighed and subsamples dried to estimate total dry mass. We recorded 22 species of vascular epiphytes, and 22 genera of cryptogams, with at least 30 species of bryophytes, liverworts and lichens. The dominant vascular epiphytes were Fascicularia bicolor (Bromeliaceae), Raukaua laetevirens, Sarmienta repens (Gesneriaceae), and filmy ferns (Hymenophyllaceae). Epiphyte loads per tree ranged between 134 and 144 kg dry mass, with 60–70% water. The hemi-epiphytic tree R. laetevirens added between 1 and 2.6 t of dry mass to each host tree. A main component of epiphyte biomass, making 70% of the weight, was detritus and roots, while leaves, stems, and fronds made up the remaining 30%. Emergent trees hold a high proportion of the regional diversity of epiphytes: 33% of all flowering epiphytes, and 50% of all filmy ferns described for Chilean temperate forests. Dry epiphyte biomass associated only with the emergent E. cordifolia trees in coastal forests was estimated in 10 t/ha. Epiphyte biomass may store up to 300 l of water in each emergent tree, and add 40–150% of photosynthetic biomass to the tree crowns. Based on this evidence, epiphytes may play key but generally neglected roles in ecosystem carbon uptake, water storage, and nutrient cycling. Moreover, emergent trees represent nuclei of biodiversity and ecosystem functions distributed throughout mature forests. Forest management should recognize large trees as significant management units for the preservation of biodiversity and ecological functions.     

Logging residue removal after thinning in Nordic boreal forests: Long-term impact on tree growth

The aim of this study was to determine the effect of whole-tree harvesting (WTH) on the growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as compared to conventional stem harvesting (CH) over 10 and 20 years. Compensatory (WTH + CoF) and normal nitrogen-based (CH + F or WTH + F) fertilisation were also studied. A series of 22 field experiments were established during 1977-1987, representing a range of site types and climatic conditions in Finland, Norway and Sweden. The treatments were performed at the time of establishment and were repeated after 10-13 years at 11 experimental sites. Seven experiments were followed for 25 years.Volume increment was on average significantly lower after WTH than after CH in both 10-year periods in the spruce stands. In the pine stands thinned only once, the WTH induced growth reduction was significant during the second 10-year period, indicating a long-term response.Volume increment of pine stands was 4 and 8% and that of spruce stands 5 and 13% lower on the WTH plots than on CH during the first and the second 10-year period, respectively. For the second 10-year period the relative volume increment of the whole-tree harvested plots tended to be negatively correlated with the amount of logging residue. Accordingly, the relative volume increment decreased more, the more logging residue was harvested, stressing the importance of developing methods for leaving the nutrient-rich needles on site.If nutrient (N, P, K) losses with the removed logging residues were compensated with fertiliser (WTH + CoF), the volume increment was equal to that in the CH plots. Nitrogen (150-180 kg ha⁻¹) or N + P fertilisation increased tree growth in all experiments except in one very productive spruce stand. Pine stands fertilised only once had a normal positive growth response during the first 10-year period, on average 13 m³ ha⁻¹, followed by a negative response of 5 m³ ha⁻¹ during the second 10-year period. The fertilisation effect of WTH + F and WTH + CoF on basal area increment was both smaller and shorter than with CH + F.     

Distribution and nest site selection of red-shouldered hawks (Buteo lineatus) in forests of northeastern Wisconsin (USA)

Red-shouldered hawks (Buteo lineatus) are threatened in Wisconsin and when nest sites are found during the cruising or marking stage of timber harvesting, the harvest is altered to accommodate the hawks. If nest site locations are known before initiation of timber harvest, foresters can employ a proactive approach to manage red-shouldered hawks while maintaining timber production. We searched for red-shouldered hawks nest sites on Marinette County Forest (MCF) which encompasses 94,000 ha in northeastern Wisconsin and is the second largest county forest in the state. We used a comparative modeling approach to evaluate distribution and habitat relationships of red-shouldered hawk nest sites in relation to a suite of environmental variables in MCF. Models were used to develop forest management recommendations for red-shouldered hawks in Wisconsin. During the spring of 2006 and 2007, we broadcasted conspecific calls to survey 1121 calling stations along forest roads and trails. We located 20 and 25 active nesting territories in 2006 and 2007, respectively (11 of which were active in both years). To understand nest site selection, we measured 22 habitat variables within 0.04-ha plots at active nest sites (n = 34) and at stratified random sites (n = 61). Logistic regression with information-theoretic model selection identified a model including greater tree species richness and closeness to forested wetland as the best-approximating model. Variable selection with Discriminant Function Analysis (DFA) indicated that nest selection was best explained by greater number of tree species, closer distance to forested wetlands, greater volume of downed woody debris, fewer small sawlogs, and increased proximity to streams. Univariate comparisons identified four of the five aforementioned variables in the DFA model as significant. Red-shouldered hawks are likely more common in Wisconsin than their state status suggests. Forest management for red-shouldered hawk nest sites should focus on increasing tree species richness, increasing down woody debris volume, and protecting forested wetlands. These recommendations may assist property managers to locate and plan for continued persistence of this species on MCF.     

Stand development patterns as a consequence of the mortality in Austrocedrus chilensis forests

The forests of Austrocedrus chilensis (D. Don) Pic. Sern. et Bizarri in Argentina suffer decline and mortality throughout their natural distribution known as ‘mal del ciprés’. While several aspects of this, to date, temporally unpredictable process of overstory tree mortality have been the focus of detailed studies, there has been little research on tree growth and stand dynamics in symptomatic forests nor stand development patterns and prediction of future stand structure. We studied 12 stands in northern Patagonia (Province of Río Negro) using stand reconstruction studies to examine the changes in stand structure over time as a consequence of overstory mortality and the implications of these structural changes on the establishment and growth of the residual overstory. Dendrochronological analyses were used to reconstruct stand establishment and structure over time, and to study past diameter growth patterns. Mortality in A. chilensis forests was variable in time among stands. As expected, overstory mortality led to the successful establishment of trees in the understory in all stands; however, the response of residual overstory trees was variable. Understory establishment was low in some stands and high in others depending on the density of the overstory. While overstory trees in almost all stands released after the onset of the mortality, the pattern was not distinctive, varying in time, number and magnitude. In some stands, growth releases occurred after single or multiple tree deaths suggesting a relationship between processes, while in others this was not the case. Even when the patterns of recruitment in the understory and the overstory response varied greatly among stands, when examined together, some general patterns emerged. This study is the first to intensively explore the dynamics of A. chilensis forests affected by ‘mal del ciprés’. Additionally, this study showed that arbitrarily categorizing disturbances as discrete or chronic masks the true process of release of growing space and the resulting stand dynamics.     

Exotic earthworms of great lakes forests: A search for indicator plant species in maple forests

The invasion of exotic earthworms in previously earthworm-free northern deciduous forests has been linked to the disappearance of forest floor litter, declines in plant species richness, and the development of monotypic stands of Carex pensylvanica. However, the impact of exotic earthworms on the regeneration of trees and understory plants is largely unknown. We examined the relationships between earthworm density, plant species richness, leaf litter accumulation, number of tree (Acer) seedlings, and cover of C. pensylvanica at 14 sites in mesic hardwood forests of the Chequamegon-Nicolet National Forest in northern Wisconsin. Earthworm biomass, especially of the genera Lumbricus, Aporrectodea, and Octolasion was negatively associated with leaf litter mass and number of Acer seedlings. Nine plant species, including two spring ephemerals (Dicentra cucullaria and Osmorhiza claytonii), a sedge (Carex deweyana), and an invasive mint (Galeopsis tetrahit) were negatively correlated with exotic earthworm densities. Dryopteris intermedia and Allium tricoccum, on the other hand, were associated with high earthworm densities. The activities of exotic earthworms appear to have significant impacts on Acer regeneration and the persistence of many herbaceous plants. Our findings suggest that the effects of exotic earthworms on litter mass are strongly related to the observed effects on species composition. Sensitive species can be used as indicators of high and low earthworm densities and might be useful for identifying forests of high conservation value where future invasions of exotic earthworms should be prevented.     

Mycorrhizas and secondary succession in aspen-conifer forests: Light limitation differentially affects a dominant early and late successional species

Plant succession and mycorrhizal fungi both play crucial roles in shaping the development of forest ecosystems. However, despite the strong potential for interactions between them, few studies have examined how patterns of forest succession affect mycorrhizal associations that a majority of plant species depend on to alleviate soil resource constraints. Fire suppression in subalpine forests over the last century has changed successional patterns in ways that may have important implications for mycorrhizal associations of forest tree species. To better understand these relationships we conducted a field and greenhouse study in which we examined mycorrhizal infection along gradients of light intensity and soil nutrient availability that develop as aspen becomes seral to conifers under longer fire return intervals. We examined whether ectomycorrhizal associations of quaking aspen (Populus tremuloides), a shade intolerant, early succession species, were more sensitive to light and soil resource limitations than subalpine fir (Abies lasiocarpa), a shade tolerant, late succession species. In the field study, ectomycorrhizal infection of aspen roots was reduced by 50% in conifer dominated stands relative to aspen stands. In contrast, subalpine fir maintained its EM associations regardless of the successional status of the stand. The greenhouse results were consistent with field results and indicated that light limitation was the driving force behind reductions in EM infection of aspen roots in later stages of succession. These results suggest that nutrient limitations constraining early successional species may be exacerbated by losses in EM associations via light limitations created by late successional species. This is one potential mechanism by which climax forest species create a competitive advantage over early successional species and these results suggest that it is likely exacerbated by longer fire return intervals.     

Long-term control of the invasive tree Ailanthus altissima: Insights from Mediterranean protected forests

Ailanthus altissima is an invasive tree species which has colonized numerous ecosystems and affected ecosystem processes worldwide. Despite its importance as an invasive species and the high economic costs incurred from its spread, there is a lack of long-term management planning for its control. Although mechanical disturbance is commonly applied, the effect that this treatment may have exhausting its resprouting ability and also its joint effect with phytochemical treatments are poorly understood, especially in Mediterranean environments. We tested three treatments (plus a control) aimed to reduce A. altissima growth in Mediterranean forests throughout 5 years of study. The treatments (one cut stump, double cut stump and cut stump with glyphosate application) were repeated annually. General plant performance (biomass, height and resprout-type density) was measured yearly during the study. Water potential and stomatal conductance were also measured at the end of the study to evaluate particular ecophysiological factors which might affect the response of A. altissima to assayed treatments, together with leaf area index. Our results show that only the cut stump with glyphosate application treatment was able to reduce the long-term growth and spread of A. altissima. The treatments applied favoured collar sprout growth in response to disturbance events (treatments) opposite to the control, where new sprouts grew mainly from the root. Treated resprouts displayed ecophysiological changes depending on the assayed treatment. To our knowledge, this is the only study testing the long-term effect of both physical disturbance and phytochemical application on A. altissima growth. Our study further refines our knowledge on the effects of repeating both commonly and newly used treatments, improving our management techniques to reduce the presence and growth of this invasive tree.