Performance of 11 tree species under different management treatments in restoration plantings in a tropical dry forest

Ecological restoration in tropical dry forests urgently needs to incorporate experimental evidence to increase effectiveness. The main barriers for tree establishment are adverse microenvironmental conditions and competition with exotic grasses. Therefore, management should address such barriers in order to enhance tree performance. We evaluated the effect of plastic mulching, grass removal, and no management on survival after 2 months and stem volume and canopy size after 2 years and integrated response index (IRI) in plantings of 11 native tree species with different growth rates in pastures near the tropical dry forest of Chamela, Mexico. Results revealed that: (1) initial seedling mortality was minimal in all treatments (8%) and lowest under no management (2%); (2) plastic mulching, but not grass removal, leads to increased size for most species, irrespective of their growth rank; (3) a trade‐off between initial plant survival and size after 2 years occurred due to plastic mulching; and (4) most species showed similar values of the IRI because of high survival, stem volume, or canopy cover. Grass removal decreased early survival of all species and increased stem volume only for one slow‐growing species. The use of plastic mulching increased stem volume for slow‐growing species, whereas fast‐growing species developed larger canopies with that treatment. Effects of grass removal and mulching seem to be very species‐specific and not dependent in growth rank of species, although overall mulching seems to provide better conditions for seedling performance than grass removal alone.     

A general ecophysiological framework for modelling the impact of pests and pathogens on forest ecosystems.

Forest insects and pathogens (FIPs) have enormous impacts on community dynamics, carbon storage and ecosystem services, however, ecosystem modelling of FIPs is limited due to their variability in severity and extent. We present a general framework for modelling FIP disturbances through their impacts on tree ecophysiology. Five pathways are identified as the basis for functional groupings: increases in leaf, stem and root turnover, and reductions in phloem and xylem transport. A simple ecophysiological model was used to explore the sensitivity of forest growth, mortality and ecosystem fluxes to varying outbreak severity. Across all pathways, low infection was associated with growth reduction but limited mortality. Moderate infection led to individual tree mortality, whereas high levels led to stand‐level die‐offs delayed over multiple years. Delayed mortality is consistent with observations and critical for capturing biophysical, biogeochemical and successional responses. This framework enables novel predictions under present and future global change scenarios.     

Underplanting degraded exotic Pinus with indigenous conifers assists forest restoration

We propose that nonharvest plantations could provide important opportunities for restoration of indigenous forest cover and related ecosystem services. We assessed the relative performance of three Podocarpaceae (podocarps) species planted into a degraded Ponderosa Pine (Pinus ponderosa) plantation, central North Island, New Zealand. We hypothesised that the degraded pine plantation overstorey could provide suitable conditions for the development of a podocarp‐dominated forest structure within ca. 50 years of underplanting and that podocarp growth would differ depending on the species suitability to the site. Rimu (Dacrydium cupressinum) significantly outperformed both Totara (Podocarpus totara) and Kahikatea (Dacrycarpus dacrydioides) in height and diameter growth. Rimu was now the structurally dominant tree where it occurred rather than pine. Per annum scaled carbon storage within Rimu stands was significantly greater than the Totara, Kahikatea or Pine stands. All podocarp species had attained a greater stand density compared to the pine overstorey. Possible reasons for the differing podocarp growth performance include different light requirements, response to soil nutrients, elevational distributions and frost susceptibility. There were significant differences in understorey species richness among the different stands of podocarp species. Underplanting accelerated successional development by incorporating late‐successional indigenous canopy dominants within the forest succession and overcame limitations imposed on forest succession at the site from its isolation from indigenous forest tree seed sources.     

A holistic approach to determine tree structural complexity based on laser scanning data and fractal analysis

The three‐dimensional forest structure affects many ecosystem functions and services provided by forests. As forests are made of trees it seems reasonable to approach their structure by investigating individual tree structure. Based on three‐dimensional point clouds from laser scanning, a newly developed holistic approach is presented that enables to calculate the box dimension as a measure of structural complexity of individual trees using fractal analysis. It was found that the box dimension of trees was significantly different among the tested species, among trees belonging to the same species but exposed to different growing conditions (at gap vs. forest interior) or to different kinds of competition (intraspecific vs. interspecific). Furthermore, it was shown that the box dimension is positively related to the trees’ growth rate. The box dimension was identified as an easy to calculate measure that integrates the effect of several external drivers of tree structure, such as competition strength and type, while simultaneously providing information on structure‐related properties, like tree growth.     

Responses of Pioneer and Later‐Successional Plant Assemblages to Created Microtopographic Variation and Soil Treatments in Riparian Forest Restoration

Riparian forest restoration generally involves introduction of later‐successional tree species, but poor species suitability to severely altered or degraded site conditions results in high mortality and poor community development. Additionally, while microtopographic heterogeneity plays a crucial role in the development of natural riparian forests, little is known regarding effects of restored or created microtopography on the development of introduced plant communities. The objective of this study was to determine the influence of created microtopography and soil treatments on early development of introduced pioneer and later‐successional plant communities in riparian forest restoration. Ridges, flats, and a mound‐and‐pool complex were created, and pioneer and later‐successional tree assemblages were planted within plots in each of these three microtopographic positions. Straw‐based erosion control mats were placed on half the plots as a source of mulch. After two growing seasons, growth and survival of the pioneer assemblage were equal among microtopographic positions, but survival of the later‐successional assemblage was significantly higher on ridges (59%) than on mounds and pools (22%) and flats (26%). A suitability index indicated that performance of the later‐successional assemblage on ridges was higher than that of the pioneer assemblage for all microtopographic positions. Flood duration explained much of the variation in plant assemblage survival, and erosion control mats had little influence on seedling survival. Restoring microtopographic features has the potential to enhance species survival and promote community development. Microtopographic restoration may be as important in riparian forest restoration as proper species selection and hydrologic reestablishment, especially at severely disturbed sites.     

Arrested succession in logging gaps: is tree seedling growth and survival limiting?

Thirty years after selective timber harvest in the Kibale National Park, Uganda, many abandoned logging gaps are dominated by Acanthus pubescens, and show little forest recovery. To examine if this arrested successional state was caused by limited tree seedling growth and survival, we planted seedlings of four forest tree species (Albizia grandibracteata, Mimusops bagshawei, Prunus africana and Uvariopsis congensis) in A. pubescens‐dominated logging gaps and in control areas of adjacent forest. To assess if clearing A. pubescens facilitates forest regeneration, we planted seedlings of two species (A. grandibracteata and U. congensis) in small clearings cut within the logging gaps. We examined mortality, growth, herbivory and site characteristics among the treatments. Finally, we described the physical attributes of the A. pubescens‐dominated gaps. Seedlings of all the four species survived and grew equally well in A. pubescens and forest treatments, and most site characteristics were also similar. Seedlings planted in clearings grew more than in either forest or A. pubescens sites. Very few established trees were found in A. pubescens sites, and most of these were near the forest edges. We also discussed the role of elephants (Loxodonta africana) and collapsing A. pubescens canopies in the maintenance of an arrested successional state in these logging gaps.     

Does Restoration Enhance Regeneration of Seasonal Deciduous Forests in Pastures in Central Brazil?

The goal of restoration is to accelerate ecosystem recovery, but in ecosystems that naturally regenerate rapidly restoration techniques need to be selected carefully to facilitate rather than impede natural recovery. We compared the effects of five restoration techniques, such as plowing the soil, removing grasses, adding forest litter, seeding, and planting nursery‐growing seedlings, on the regeneration of seasonal deciduous forest trees in four abandoned pastures in central Brazil. We monitored all woody stems immediately prior to treatments and again 14 months after the treatments. We recorded an average of 16,663 tree stems per hectare and a total of 83 species before implementing treatments. Planting strongly increased species richness; adding litter and seeding had weaker positive effects on richness; and plowing and grass removal had no effect. Plowing substantially reduced the density of naturally established stems. Despite the high survival of planted seedlings, stem density in planting treatments did not change because the tractor and digging holes to plant seedlings caused mortality of naturally regenerating seedlings. Tree stems grew more in the grass release plots than in the control plots. Our results suggest that early succession of seasonal deciduous forest in pastures in the region studied does not need to be stimulated once the perturbation is stopped and that intensive restoration efforts may actually slow recovery. We recommend only enrichment planting of seedlings that are not able to resprout.     

Drought's legacy: multiyear hydraulic deterioration underlies widespread aspen forest die‐off and portends increased future risk

Forest mortality constitutes a major uncertainty in projections of climate impacts on terrestrial ecosystems and carbon‐cycle feedbacks. Recent drought‐induced, widespread forest die‐offs highlight that climate change could accelerate forest mortality with its diverse and potentially severe consequences for the global carbon cycle, ecosystem services, and biodiversity. How trees die during drought over multiple years remains largely unknown and precludes mechanistic modeling and prediction of forest die‐off with climate change. Here, we examine the physiological basis of a recent multiyear widespread die‐off of trembling aspen (Populus tremuloides) across much of western North America. Using observations from both native trees while they are dying and a rainfall exclusion experiment on mature trees, we measure hydraulic performance over multiple seasons and years and assess pathways of accumulated hydraulic damage. We test whether accumulated hydraulic damage can predict the probability of tree survival over 2 years. We find that hydraulic damage persisted and increased in dying trees over multiple years and exhibited few signs of repair. This accumulated hydraulic deterioration is largely mediated by increased vulnerability to cavitation, a process known as cavitation fatigue. Furthermore, this hydraulic damage predicts the probability of interyear stem mortality. Contrary to the expectation that surviving trees have weathered severe drought, the hydraulic deterioration demonstrated here reveals that surviving regions of these forests are actually more vulnerable to future droughts due to accumulated xylem damage. As the most widespread tree species in North America, increasing vulnerability to drought in these forests has important ramifications for ecosystem stability, biodiversity, and ecosystem carbon balance. Our results provide a foundation for incorporating accumulated drought impacts into climate–vegetation models. Finally, our findings highlight the critical role of drought stress accumulation and repair of stress‐induced damage for avoiding plant mortality, presenting a dynamic and contingent framework for drought impacts on forest ecosystems.     

Species‐Specific Barriers to Tree Regeneration in High Elevation Habitats of West Virginia

Herbaceous competition and herbivory have been identified as critical barriers to restoration of native tree species in degraded landscapes around the world; however, the combined effects of competition and herbivory are poorly understood. We experimentally manipulated levels of herbivory and herbaceous competition and analyzed the response of tree seedling performance over three growing seasons as a function of species and habitat in north‐central West Virginia. Four native tree species were planted in old field and forest experimental plots: Castanea dentata (American chestnut), Quercus rubra (red oak), Acer saccharum (sugar maple), and Picea rubens (red spruce). Red spruce demonstrated the highest growth increment and greatest survival (64%) and most consistent results among treatments and habitats. Red spruce survival was not reduced in the presence of Odocoileus virginianus (white‐tailed deer) browse and herbaceous competition; however, growth was improved by suppression of herbaceous competition. We suspect that this deciduous forest landscape would regenerate to a red spruce dominated forest if seed source was available. In contrast, the other three species tested had very low survival when exposed to deer and were more responsive to competing vegetation and habitat type. American chestnut had low survival and growth across all treatments, suggesting basic climate limitations. Vigorous natural regeneration of Prunus serotina (black cherry) occurred in forest plots where both competing herbs and deer were excluded. Our results demonstrated the importance of testing multiple potential recruitment barriers and species at once and the need for species and habitat‐specific restoration treatments.     

Demographic costs and benefits of natural regeneration during tropical forest restoration

For tropical forest restoration to result in long‐term biodiversity gains, native trees must establish self‐sustaining populations in degraded sites. While many have asked how seedling recruitment varies between restoration treatments, the long‐term fate of these recruits remains unknown. We address this research gap by tracking natural recruits of 27 species during the first 7 years of a tropical forest restoration experiment that included both planted and naturally regenerating plots. We used an individual‐based model to estimate the probability that a seedling achieves reproductive maturity after several years of growth and survival. We found an advantage for recruits in naturally regenerating plots, with up to 40% increased probability of reproduction in this treatment, relative to planted plots. The demographic advantage of natural regeneration was highest for mid‐successional species, with relatively minor differences between treatments for early‐successional species. Our research demonstrates the consequences of restoration decision making across the life cycle of tropical tree species.     

Non‐native fruit trees facilitate colonization of native forest on abandoned farmland

Ecological restoration of abandoned, formerly forested farmland can improve the delivery of ecosystem services and benefit biodiversity conservation. Restoration programs can involve removing isolated, non‐native trees planted by farmers for fruit or wood. As such “legacy” trees can attract seed dispersers and create microclimates that help native seedlings to establish, removing them may actually slow forest recovery. Working on abandoned farmland in Kibale National Park, Uganda, we evaluated the effect of legacy trees on forest recovery by measuring the number, diversity, and biomass of native seedlings and saplings regenerating in plots centered on avocado (Persea americana), mango (Mangifera indica), and Eucalyptus legacy trees compared with adjacent plots without legacy trees. The assemblages of native, forest‐dependent tree species in plots around avocado and mango trees were distinct from each other and from those around eucalyptus and all the near‐legacy plots. In particular, avocado plots had higher stem density and species richness of forest‐dependent species than near‐avocado plots, particularly large‐seeded, shade‐tolerant, and animal‐dispersed species—key targets of many restoration plans. Furthermore, many of the species found in high numbers were among those failing to establish in ongoing large‐scale forest restoration in Kibale. Taken together, our results demonstrate that the legacy trees facilitate the dispersal and establishment of native tree species. Retaining the existing legacy trees for a number of years could usefully complement existing management strategies to restore more biodiverse native forest in degraded lands. However, careful monitoring is needed to ensure that the legacy trees do not themselves establish.     

Restoration of Old Forest Features in Coast Redwood Forests Using Early‐stage Variable‐density Thinning

To accelerate development of old forest features in coast redwood, two thinning treatments and an unthinned control were compared in three treatment areas in north coastal California. One thinning treatment was designed to restore old forest densities of 125 trees/ha and the other 250 trees/ha representing a one‐step and partial treatments to the desired stand density. Four years after treatment, numbers of trees had increased in the thinning treatments due to recruitment of new trees, but had decreased in the control due to self‐thinning. Residual trees increased in stem volume following thinning by 128% in low‐density thinning compared to 70% in the controls indicating thinning accelerated stand development. The thinning treatments also moved the species composition of these stands to a greater proportion of redwood. Considerable slash was produced by the thinning treatments but was decomposing rapidly. Black bears damaged approximately 15% of all trees and more than 38% of residual trees in the thinned treatments compared to less than 2% of all trees in the control. This damage included killing some trees and damaging other trees that survived. Decisions over restoration densities in these stands are complicated by prolonged stand development, and balancing risks and costs. In this case, the bears represent a stochastic factor that dramatically increases risk. Thinning appears to be an effective means of enhancing old forest development by accelerating tree growth, modifying species composition, and increasing stand‐level variability. Continued monitoring will be necessary to evaluate long‐term trends in density relative to effects of bear damage.     

Tropical soils degraded by slash‐and‐burn cultivation can be recultivated when amended with ashes and compost

In many tropical regions, slash‐and‐burn agriculture is considered as a driver of deforestation; the forest is converted into agricultural land by cutting and burning the trees. However, the fields are abandoned after few years because of yield decrease and weed invasion. Consequently, new surfaces are regularly cleared from the primary forest. We propose a reclamation strategy for abandoned fields allowing and sustaining re‐cultivation. In the dry region of south‐western Madagascar, we tested, according to a split‐plot design, an alternative selective slash‐and‐burn cultivation technique coupled with compost amendment on 30–year‐old abandoned fields. Corn plants (Zea mays L.) were grown on four different types of soil amendments: no amendment (control), compost, ashes (as in traditional slash‐and‐burn cultivation), and compost + ashes additions. Furthermore, two tree cover treatments were applied: 0% tree cover (as in traditional slash‐and‐burn cultivation) and 50% tree cover (selective slash‐and‐burn). Both corn growth and soil fertility parameters were monitored during the growing season 2015 up to final harvest. The amendment compost + ashes strongly increased corn yield, which was multiplied by 4–5 in comparison with ashes or compost alone, reaching 1.5 t/ha compared to 0.25 and 0.35 t/ha for ashes and compost, respectively. On control plots, yield was negligible as expected on these degraded soils. Structural equation modeling evidenced that compost and ashes were complementary fertilizing pathways promoting soil fertility through positive effects on soil moisture, pH, organic matter, and microbial activity. Concerning the tree cover treatment, yield was reduced on shaded plots (50% tree cover) compared to sunny plots (0% tree cover) for all soil amendments, except ashes. To conclude, our results provide empirical evidence on the potential of recultivating tropical degraded soils with compost and ashes. This would help mitigating deforestation of the primary forest by increasing lifespan of agricultural lands.     

Does Tree Species Composition Affect Productivity in a Tropical Planted Forest?

With growing pressure on primary forests from destructive land uses, increasing the diversity of native species plantations can increase ecosystem service provision, such as timber production or carbon sequestration, thus better supporting sustainable livelihoods. Understanding the effects of tree species composition on productivity can inform plantation design and ecological restoration strategies. However, tree species composition effects have been neglected in experimental biodiversity‐ecosystem function (BEF) research. This study uses a 10‐yr data set from one of the first tropical planted forest experiments established with native species and designed for BEF research at scales relevant to forest management. At our site in Sardinilla, Panama, we established plots containing 6 species from a pool of 18, in four combinations, to investigate how composition affects species and plot productivity. We used basal area as a proxy for productivity through time, measured annually, and summed this at species and plot levels for analysis. We found that plots that differed in species composition appeared to differ in temporal rate of basal area increase, but did not differ in BA after 10 yr. Species were generally consistent in size between compositions, and composition performance was correlated with the size of component species, suggesting that species identities were most important in determining plot productivity. Our results suggest that species choice can be based on preferences for individual species, as species performance was consistent across composition contexts. We make recommendations for the use of particularly productive species that also provide multiple services such as Guazuma ulmifolia, Spondias mombin, and Anacardium excelsum.     

Demographic controls of aboveground forest biomass across North America

Ecologists have limited understanding of how geographic variation in forest biomass arises from differences in growth and mortality at continental to global scales. Using forest inventories from across North America, we partitioned continental‐scale variation in biomass growth and mortality rates of 49 tree species groups into (1) species‐independent spatial effects and (2) inherent differences in demographic performance among species. Spatial factors that were separable from species composition explained 83% and 51% of the respective variation in growth and mortality. Moderate additional variation in mortality (26%) was attributable to differences in species composition. Age‐dependent biomass models showed that variation in forest biomass can be explained primarily by spatial gradients in growth that were unrelated to species composition. Species‐dependent patterns of mortality explained additional variation in biomass, with forests supporting less biomass when dominated by species that are highly susceptible to competition (e.g. Populus spp.) or to biotic disturbances (e.g. Abies balsamea).     

Frequent,low‐amplitude disturbances drive high tree turnover rates on a remote,cyclone‐prone Polynesian island

Aim How important are frequent, low‐intensity disturbances to tree community dynamics of a cyclone‐prone forest? We tested the following hypotheses concerning the ‘inter‐cataclysm’ period on a remote Polynesian island: (1) tree turnover would be high and recruitment rates would be significantly higher than mortality; (2) low‐intensity disturbance would result in a marginal increase in tree mortality in the short term; (3) turnover would vary among species and would be associated with plant traits linked to differences in life history; and (4) mortality and recruitment events would be spatially non‐random. Location Tutuila, a volcanic island in the Samoan Archipelago, Polynesia. Methods We censused the tree (stem diameter ≥ 10 cm) community in 3.9 ha of tropical forest three times over a 10‐year period, 1998–2008. We calculated annual mortality, recruitment and turnover rates for 36 tree species. We tested for non‐random spatial patterns and predictors of mortality, and non‐random spatial patterns of tree recruitment. A 2004 cyclone passing within 400 km allowed us to measure the effects of a non‐cataclysmic disturbance on vital rates. Results Annual turnover was 2.8% and annual recruitment was 3.6%; these are some of the highest rates in the tropics, and likely to be a response to a cyclone that passed < 50 km from Tutuila in 1991. Species turnover rates over 10 years were negatively correlated with wood specific gravity, and positively correlated with annual stem diameter increment. Mortality was spatially aggregated, and a function of site, species and an individual’s growth rate. Recruitment was highest on ground with low slope. The low‐magnitude cyclone disturbance in 2004 defoliated 29% of all trees, but killed only 1.8% of trees immediately and increased annual mortality over 5 years by 0.7%. Main conclusions The inter‐cataclysm period on Tutuila is characterized by frequent, low‐amplitude disturbances that promote high rates of tree recruitment and create a dynamic, non‐equilibrium or disturbed island disequilibrium tree community. Species with low wood density and fast growth rates have enhanced opportunities for recruitment between cataclysms, but also higher probabilities of dying. Our results suggest that increases in the frequency of cyclone activity could shift relative abundances towards disturbance‐specialist species and new forest turnover rates.     

Geographical distributions in tropical trees: can geographical range predict performance and habitat association in co‐occurring tree species?

Aim A major floristic and climatic transition from aseasonal to seasonal evergreen tropical forest (the Kangar–Pattani Line; KPL) exists in the Indo‐Sundaic region of Southeast Asia. Mechanisms constraining species distribution here are at present poorly understood, but it is hypothesized that species differ in their tolerances of abiotic factors, in particular water availability. Under this hypothesis, we anticipate differences in performance or habitat preferences, or both, of species differing in distribution with respect to the KPL. The aim of this study is to test whether geographical distributions can be used to explain variation in growth, mortality and habitat preferences in co‐occurring tree species differing in their distribution in relation to the KPL. Location Pasoh Forest Reserve, Negeri Sembilan, Malaysia; south of the KPL. Methods All tree species within a 50‐ha forest dynamics plot were classified as widespread or southern based upon their distributions in relation to the KPL and as habitat specialists or generalists based on spatial association with soil‐based habitat categories. Growth and mortality rates, variation in growth and mortality with respect to soil type, and levels of habitat association were quantified for species with different geographical distributions. Results Differences existed in species performance based upon geographical distributions. Specifically, widespread species had lower growth rates than did species restricted to the aseasonal forests. Mortality rates did not differ as a function of geographical distribution. The growth responses of species to soil habitats also diverged, such that differences in performance of widespread species among soil types were more conservative than those of species restricted in their distribution to the aseasonal forests. However, the proportion of species showing positive habitat associations did not differ significantly between widespread and southern species. Main conclusions Distribution‐based differences in species performance and response to soil type support the hypothesis that species tolerant of wider climatic variation perform less well in any given environment due to limitations on plasticity. These performance differences provide quantitative evidence of the role of climatic transitions in determining tree species distributions in relation to the Kangar–Pattani Line in the Indo‐Malay region. Such differences in performance have important implications for our understanding of biodiversity gradients and responses of Indo‐Sundaic forests to climate change.     

Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Impacts of the emerald ash borer (Agrilus planipennis Fairmaire) induced ash (Fraxinus spp.) mortality on forest carbon cycling and successional dynamics in the eastern United States

Invasive species are widely recognized as altering species and community dynamics, but their impacts on biogeochemical cycling and ecosystem processes are less understood. The emerald ash borer (Agrilus planipennis Fairmaire) is a phloem feeding beetle that was inadvertently introduced to the US in the 1990s and relies solely on ash trees (Fraxinus spp.) to complete its life cycle. Ash trees have a wide geographic distribution and are an important component of many different forest types in the US. The larval feeding behavior of the emerald ash borer (EAB) effectively girdles the tree’s phloem tissue resulting in tree mortality in as little as 2 years and stand mortality in as little as 5 years. Using the forest inventory and analysis database, we found that forest lands in the lower 48 states hold approximately 8.7 billion ash trees and saplings, which represent ~2.5 % of the aboveground forest carbon mass. Furthermore, we measured tree growth in 7 EAB impacted and 5 non-EAB impacted temperate forests in the Midwestern United States to quantify the impacts of EAB induced tree mortality on tree growth. We hypothesized that the initial C lost would be partly compensated for by the enhanced non-ash tree growth in EAB-impacted regions relative to non-EAB impacted regions. The EAB disturbance enhanced growth of non-ash trees in the EAB impacted region relative to the non-EAB impacted region. Results also indicate that in EAB impacted areas, growth of trees from the genera Acer and Ulmus responded most positively. Finally, we quantified annual biometric net primary productivity of the EAB impacted forests and compared these quantities to modeled growth of these forests in the absence of EAB and found that large scale ash tree mortality has reduced short term regional forest productivity. The loss of ash biometric net primary productivity is, in part compensated by enhanced growth of non-ash species. As expected, EAB disturbance severity was greater in forests with higher basal areas of ash. This study illustrates the ecosystem and regional scale impacts of invasive pest-induced disturbance on biogeochemical cycling and forest species composition.