Sustaining eastern oak forests: Synergistic effects of fire and topography on vegetation and fuels.

Across much of the eastern United States, oak forests are undergoing mesophication as shade-tolerant competitors become more abundant and suppress oak regeneration. Given the historical role of anthropogenic surface fires in promoting oak dominance, prescribed fire has become important in efforts to reverse mesophication and sustain oaks. In 2000 we established the Ohio Hills Fire and Fire Surrogate (FFS) study to examine whether repeated prescribed fire (Fire), mechanical partial harvest (Mech), and their combined application (Mech + Fire) reduced the dominance of subcanopy mesophytic competitors, increased the abundance of large oak-hickory advance regeneration, created a more diverse and productive ground-layer flora, and produced fuel beds more conducive to prescribed fire, reducing the risk of high-severity wildfire. Here we report on the ~20-year effects of treatments on vegetation and fuels and examine the support for interactive effects across a topographic-moisture and energy gradient. In general, we found that Fire and Mech + Fire treatments tended to reverse mesophication while the Mech-only treatment did not. The moderate and occasionally high-intensity fires resulted in effects that were ultimately very similar between the two fire treatments but were modulated by topography with increasing fire severity on drier sites. In particular, we found support for an interaction effect between treatment and topography on forest structure and tree regeneration responses. Fire generally reduced mesophytic tree density in the midstory and sapling strata across all site conditions, while leading to substantial gains in the abundance of large oak-hickory advance regeneration on dry and intermediate landscape positions. Fire also promoted ground-layer diversity and created compositionally distinct communities across all site conditions, primarily through the increased richness of native perennial herbs. However, the fire had limited effects on fine surface fuel loading and increased the loading of large woody fuels, potentially increasing the risk of high-severity wildfire during drought conditions. We conclude that two decades of repeated fires, with and without mechanical density reduction, significantly shifted the trajectory of mesophication across most of the landscape, particularly on dry and intermediate sites, highlighting the capacity of a periodic fire regime to sustain eastern oak forests and promote plant diversity but modulated by topography.

Stasis in forest regeneration following deer exclusion and understory gap creation: A 10-year experiment.

Chronically elevated ungulate browse pressure in temperate forests worldwide often generates ecological legacies characterized by low plant diversity and contributes to the formation of dense, nearly monodominant, and highly recalcitrant layers of understory vegetation. Once established, these recalcitrant layers combined with continued browsing may jointly constrain tree establishment and diversity so completely that understory recovery may be unattainable without mitigating browse pressure, the recalcitrant layer, or both. Here, we investigate the independent and synergistic effects of both white-tailed deer (Odocoileus virginianus) browsing and hay-scented fern (Dennstaedtia punctilobula) competition on tree regeneration in a 10-year experiment. Specifically, we examine how tree seedling establishment, growth, and composition are filtered by fern cover versus fern removal (gaps), browser presence versus absence (exclosures), and their combined effects during 10 years at three hardwood forest sites in Pennsylvania, USA. Fern gaps enhanced establishment for multiple tree species, increasing seedling density and diversity, particularly in the first 3 years post-treatment, and enhancing richness (≤1 species) over the course of the experiment. Excluding deer for a decade increased the height growth of other regeneration and altered species composition, but had no effect on diversity, richness, and density. Notably, we observed higher Prunus serotina seedling densities outside exclosures, possibly due to greater secondary dispersal. We argue that browsing legacies in second growth forests established at the turn of the last century created two conditions inimical to diverse forest regeneration: an overstory dominated by two species, P. serotina and Acer rubrum (86% of basal area), and a dense recalcitrant understory layer dominated by a native fern. The first condition limits propagule supply, the second strongly filters seedling establishment, and both create impoverished forest understories composed of few individuals and species. In undisturbed forest understories, the inertia toward impoverishment was sustained across the decade even where browsing was eliminated. Consequently, stand replacing disturbances (whether natural or anthropogenic) that disrupt the understory layer and reinitiate succession may be necessary to propel forests out of their current stasis and down a pathway leading to greater diversity.

Phytochemicals Involved in Plant Resistance to Leporids and Cervids: a Systematic Review.

Non-nutritive phytochemicals (secondary metabolites and fibre) can influence plant resistance to herbivores and have ecological impacts on animal and plant population dynamics. A major hindrance to the ecological study of these phytochemicals is the uncertainty in the compounds one should measure, especially when limited by cost and expertise. With the underlying goal of identifying proxies of plant resistance to herbivores, we performed a systematic review of the effects of non-nutritive phytochemicals on consumption by leporids (rabbits and hares) and cervids (deer family). We identified 133 out of 1790 articles that fit our selection criteria (leporids = 33, cervids = 97, both herbivore types = 3). These articles cover 18 species of herbivores, on four continents. The most prevalent group of phytochemicals in the selected articles was phenolics, followed by terpenes for leporids and by fibre for cervids. In general, the results were variable but phenolic concentration seems linked with high resistance to both types of herbivores. Terpene concentration is also linked to high plant resistance; this relationship seems driven by total terpene content for cervids and specific terpenes for leporids. Tannins and fibre did not have a consistent positive effect on plant resistance. Because of the high variability in results reported and the synergistic effects of phytochemicals, we propose that the choice of chemical analyses must be tightly tailored to research objectives. While researchers pursuing ecological or evolutionary objectives should consider multiple specific analyses, researchers in applied studies could focus on a fewer number of specific analyses. An improved consideration of plant defence, based on meaningful chemical analyses, could improve studies of plant resistance and allow us to predict novel or changing plant-herbivore interactions.

A regional assessment of white-tailed deer effects on plant invasion.

Herbivores can profoundly influence plant species assembly, including plant invasion, and resulting community composition. Population increases of native herbivores, e.g. white-tailed deer (Odocoileus virginianus), combined with burgeoning plant invasions raise concerns for native plant diversity and forest regeneration. While individual researchers typically test for the impact of deer on plant invasion at a few sites, the overarching influence of deer on plant invasion across regional scales is unclear. We tested the effects of deer on the abundance and diversity of introduced and native herbaceous and woody plants across 23 white-tailed deer research sites distributed across the east-central and north-eastern USA and representing a wide range of deer densities and invasive plant abundance and identity. Deer access/exclusion or deer population density did not affect introduced plant richness or community-level abundance. Native and total plant species richness, abundance (cover and stem density) and Shannon diversity were lower in deer-access vs. deer-exclusion plots. Among deer-access plots, native species richness, native and total cover, and Shannon diversity (cover) declined as deer density increased. Deer access increased the proportion of introduced species cover (but not of species richness or stem density). As deer density increased, the proportion of introduced species richness, cover and stem density all increased. Because absolute abundance of introduced plants was unaffected by deer, the increase in proportion of introduced plant abundance is likely an indirect effect of deer reducing native cover. Indicator species analysis revealed that deer access favoured three introduced plant species, including Alliaria petiolata and Microstegium vimineum, as well as four native plant species. In contrast, deer exclusion favoured three introduced plant species, including Lonicera japonica and Rosa multiflora, and 15 native plant species. Overall, native deer reduced community diversity, lowering native plant richness and abundance, and benefited certain invasive plants, suggesting pervasive impacts of this keystone herbivore on plant community composition and ecosystem services in native forests across broad swathes of the eastern USA.

Unearthing the hidden world of roots: Root biomass and architecture differ among species within the same guild.

The potential benefits of planting trees have generated significant interest with respect to sequestering carbon and restoring other forest based ecosystem services. Reliable estimates of carbon stocks are pivotal for understanding the global carbon balance and for promoting initiatives to mitigate CO2 emissions through forest management. There are numerous studies employing allometric regression models that convert inventory into aboveground biomass (AGB) and carbon (C). Yet the majority of allometric regression models do not consider the root system nor do these equations provide detail on the architecture and shape of different species. The root system is a vital piece toward understanding the hidden form and function roots play in carbon accumulation, nutrient and plant water uptake, and groundwater infiltration. Work that estimates C in forests as well as models that are used to better understand the hydrologic function of trees need better characterization of tree roots. We harvested 40 trees of six different species, including their roots down to 2 mm in diameter and created species-specific and multi-species models to calculate aboveground (AGB), coarse root belowground biomass (BGB), and total biomass (TB). We also explore the relationship between crown structure and root structure. We found that BGB contributes ~27.6% of a tree's TB, lateral roots extend over 1.25 times the distance of crown extent, root allocation patterns varied among species, and that AGB is a strong predictor of TB. These findings highlight the potential importance of including the root system in C estimates and lend important insights into the function roots play in water cycling.

Evaluating the ecological impacts of salvage logging: can natural and anthropogenic disturbances promote coexistence?

Salvage logging following windthrow is common throughout forests worldwide even though the practice is often considered inimical to forest recovery. Because salvaging removes trees, crushes seedlings, and compacts soils, many warn this practice may delay succession, suppress diversity, and alter composition. Here, over 8 yr following windthrow, we experimentally evaluate how salvaging affects tree succession across 11 gaps in Eastern deciduous forests of Pennsylvania, wherein each gap was divided into salvaged and control (unsalvaged) halves. Our gaps vary in size and windthrow severity, and we explicitly account for this variation as well as variation in soil disturbance (i.e., scarification) resulting from salvaging so that our results would be generalizable. Salvage logging had modest and ephemeral impacts on tree succession. Seedling richness and density declined similarly over time in both salvaged and unsalvaged areas as individuals grew into saplings. The primary impact of salvaging on succession occurred where salvaging scarified soils. Here, salvaging caused 41 to 82% declines in sapling abundance, richness, and diversity, but these differences largely disappeared within 5 yr. Additionally, we documented interactions between windthrow severity and scarification. Specifically, low-severity windthrow and scarification combined reinforced dominance by shade-tolerant and browse-tolerant species (Acer pensylvanicum, Fagus grandifolia). In contrast, high windthrow severity and scarification together reduced the density of a fast-growing pioneer tree (Prunus pensylvanica) and non-tree vegetation cover by 75% and 26%, respectively. This reduction enhanced the recruitment of two mid-successional tree species, Acer rubrum and Prunus serotina, by 2 and 3-fold, respectively. Thus, our findings demonstrate that salvaging creates novel microsites and mitigates competing vegetation, thereby enhancing establishment of important hardwoods and promoting tree species coexistence. Our results, coupled with an assessment of 27 published post-windthrow salvage studies, suggest short-term studies may overestimate the impact of salvaging on regeneration. We conclude that the ecological costs and benefits of salvaging depend upon the variation in canopy and soil disturbance severity as well as the timescale at which effects are evaluated. Thus, our findings are inconsistent with the view that salvaging inexorably undermines plant diversity; rather we suggest salvaging can promote tree species coexistence within various contexts.

Pervasive interactions between ungulate browsers and disturbance regimes promote temperate forest herbaceous diversity.

Disruptions to historic disturbance and herbivory regimes have altered plant assemblages in forests worldwide. An emerging consensus suggests that these disruptions often result in impoverished forest biotas. This is particularly true for eastern U.S. deciduous forests where large gaps and understory fires were once relatively common and browsers were far less abundant. Although much research has focused on how disturbance and browsers affect tree diversity, far less attention has been devoted to forest understories where the vast majority (>75%) of the vascular species reside. Here we test the hypothesis that the reintroduction of disturbances resembling historic disturbance regimes and moderate levels of ungulate browsing enhance plant diversity. We explore whether once-common disturbances and their interaction with the top-down influence of browsers can create conditions favorable for the maintenance of a rich herbaceous layer in a region recognized as a temperate biodiversity hotspot in West Virginia, U.S.A. We tested this hypothesis via a factorial experiment whereby we manipulated canopy gaps (presence/absence) of a size typically found in old-growth stands, low-intensity understory fire (burned/unburned), and deer browsing (fenced/unfenced). We tracked the abundance and diversity of more than 140 herb species for six years. Interactions among our treatments were pervasive. The combination of canopy gaps and understory fire increased herbaceous layer richness, cover, and diversity well beyond either disturbance alone. Furthermore, we documented evidence that deer at moderate levels of abundance promote herbaceous richness and abundance by preferentially browsing fast-growing pioneer species that thrive following co-occurring disturbances (i.e., fire and gaps). This finding sharply contrasts with the negative impact browsers have when their populations reach levels well beyond those that occurred for centuries. Although speculative, our results suggest that interactions among fire, canopy gaps, and browsing provided a variable set of habitats and conditions across the landscape that was potentially capable of maintaining much of the plant diversity found in temperate forests.

The herb community of a tropical forest in central Panamá: dynamics and impact of mammalian herbivores.

Mammals are hypothesized to either promote plant diversity by preventing competitive exclusion or limit diversity by reducing the abundance of sensitive plant species through their activities as browsers or disturbance agents. Previous studies of herbivore impacts in plant communities have focused on tree species and ignored the herbaceous community. In an experiment in mature-phase, tropical moist forest sites in central Panamá, we studied the impact of excluding ground-dwelling mammals on the richness and abundance of herbs in 16, 30x45-m plots. Within each plot, we censused the herbaceous community in 28, 2x2-m subplots (1,792 m2 total area sampled). We identified over 54 species of herbs averaging 1.21 ramets m-2 and covering approximately 4.25% of the forest floor. Excluding mammals for 5 years had no impact on overall species richness. Within exclosures, however, there was a significant two-fold increase in the density of rare species. Overall herbaceous density and percent cover did not differ between exclosures and adjacent control plots, although cover did increase over time. Mammalian exclusion significantly increased the total cover of three-dominant herb species, Pharus latifolius, Calathea inocephala, and Adiantum lucidum, but did not affect their density. This study represents one of the most extensive herbaceous community censuses conducted in tropical forests and is among a few that quantify herbaceous distribution and abundance in terms of both density and cover. Additionally, this work represents the first community level test of mammalian impacts on the herbaceous community in a tropical forest to date. Our results suggest that ground dwelling mammals do not play a key role in altering the relative abundance patterns of tropical herbs in the short term. Furthermore, our results contrast sharply with prior studies on similar temporal and spatial scales that demonstrate mammals strongly alter tree seedling composition and reduce seedling density. Thus, we question the pervasiveness of top-down control on tropical plant communities and the paradigm that defaunation will inexorably lead to widespread, catastrophic shifts in plant communities.