Influence of white-tailed deer on nitrogen cycling and vegetative community change in canopy gaps in a hemlock-northern hardwood forest

Our research explored the influence of deer and gap size on nitrogen cycling, soil compaction, and vegetation trajectories in twelve canopy gaps of varying sizes in a hemlock-northern hardwood forest. Each gap contained two fenced and two unfenced plots. Gap size, soil compaction, winter deer use, and available nitrogen were measured in 2011. Vegetation was assessed in 2007 and 2011, and non-metric multi-dimensional scaling was used to determine vegetative change. Results show that winter deer use was greater in smaller gaps. Deer accessibility did not influence compaction but did significantly increase total available nitrogen in April. April ammonium, April nitrate, and May nitrate were positively related to gap size. The relationship between gap size and vegetative community change was positive for fenced plots but unrelated for unfenced plots. In conclusion, deer are positively contributing to nitrogen dynamics and altering the relationship between canopy gap size and vegetative community change.

Canopy gaps promote selective stem-cutting by small mammals of two dominant tree species in an African lowland forest: the importance of seedling chemistry

Small mammals can impede tree regeneration by injuring seedlings and saplings in several ways. One fatal way is by severing their stems, but apparently this type of predation is not well-studied in tropical rain forest. Here, we report on the incidence of 'stem-cutting' to new, wild seedlings of two locally dominant, canopy tree species monitored in 40 paired forest understorey and gap-habitat areas in Korup, Cameroon following a 2007 masting event. In gap areas, which are required for the upward growth and sapling recruitment of both species, 137 seedlings of the long-lived, light-demanding, fast-growing large tropical tree (Microberlinia bisulcata) were highly susceptible to stem-cutting (83% of deaths) - it killed 39% of all seedlings over a c. 2-y period. In stark contrast, seedlings of the more shade-tolerant, slower-growing tree species (Tetraberlinia bifoliolata) were hardly attacked (4.3%). In the understorey, however, stem-cutting was virtually absent. Across the gap areas, the incidence of stem-cutting of M. bisulcata seedlings showed significant spatial variation that could not be explained significantly by either canopy openness or Janzen-Connell type effects (proximity and basal area of conspecific adult trees). To examine physical and chemical traits that might explain the species difference to being cut, bark and wood tissues were collected from a separate sample of seedlings in gaps (i.e. not monitored for stem-cutting). These analyses suggested that, compared with T. bifoliolata, the lower stem density, higher Mg and K and fatty acid concentrations in bark, and fewer phenolic and terpene compounds in M. bisulcata seedlings made them more palatable and attractive to small-mammal predators, likely rodents. We conclude that selective stem-cutting is a potent countervailing force to the current local canopy dominance of the grove-forming M. bisulcata by limiting the recruitment and abundance of its saplings. Given the ubiquity of gaps and ground-dwelling rodents in pantropical forests, it would be surprising if this form of lethal browsing was restricted to Korup.

The successional dynamics of lightning-initiated canopy gaps in the mangrove forests of Shark River, Everglades National Park, USA

Gap succession is a significant determinant of structure and development in most forest communities. Lightning strikes are an important source of canopy gaps in the mangrove forest of Everglades National Park. I investigated the successional dynamics of lightning-initiated canopy gaps to determine their influence on forest stand structure of the mixed mangrove forests ( Rhizophora mangle, Laguncularia racemosa, and Avicennia germinans ) of the Shark River. I measured gap size, gap shape, light environment, soil characteristics, woody debris, and fiddler crab abundance. I additionally measured the vegetative composition in a chronosequences of gap successional stages (new, recruiting, and growing gaps). I recorded survivorship, recruitment, growth and soil elevation dynamics within a subset of new and growing gaps. I determined the relationship between intact forest soil elevation and site hydrology in order to interpret the effects of lightning disturbance on soil elevation dynamics. ^ Gap size averaged 289 ± 20 m2 (± 1SE) and light transmittance decreased exponentially as gaps filled with saplings. Fine woody debris was highest in recruiting gaps. Soil strength was lower in the gaps than in the forest. The abundance of large and medium fiddler crab burrows increased linearly with total seedling abundance. Soil surface elevation declined in newly formed lightning gaps; this loss was due to a combination of superficial erosion (8.5 mm) and subsidence (60.9 mm). A distinct two-cohort recruitment pattern was evident in the seedling/sapling surveys, suggesting a partitioning of the succession between individuals present before and after lightning strike. In new gaps, the seedling recruitment rate was twice as high as in forest and the sapling population increased. At the growing gap stage, R. mangle seedling mortality was 10 times greater and sapling mortality was 13 times greater than recruitment. Growing gaps had reduced seedling stem elongation, sapling growth and adult growth. However, a few individuals (R. mangle saplings) were able to recruit into the adult life stage. In conclusion, the high density of R. mangle seedlings and saplings imply that lightning strike disturbances in these mangrove forests favor their recruitment over that of A. germinans and L. racemosa. ^

Canopy cover mediates interactions between a specialist caterpillar and seedlings of a neotropical tree

1 Light availability may be crucial for understanding dynamics of plant–herbivore interactions in temperate and tropical forest communities. This is because local light availability can influence both tree seedling tolerance and susceptibility to herbivory – yet how they mediate levels of insect herbivory that vary with the density of host population is virtually unknown. Here we tested predictions of three key, non-mutually exclusive hypotheses of plant–herbivore interactions: the Limiting Resource Model (LRM), the Plant Vigour Hypothesis (PVH), and the Janzen-Connell Mechanism (JCM). 2 In an Amazonian forest, we planted Swietenia macrophylla seedlings (c. 5 months old) into natural canopy gaps and the shaded understorey and simulated the damage patterns of the specialist herbivore moth, Steniscadia poliophaea, by clipping seedling leaves. Over the next 8 months, we monitored seedling performance in terms of growth and survivorship and also quantified herbivory to new young leaves on a seasonal basis. 3 In support of the LRM, severe leaf damage (≥ 50%) was lethal for Swietenia macrophylla seedlings in the understorey, but in gaps only reduced seedling growth. In support of the PVH, gap seedlings suffered greater post-simulated herbivory (up to 100% defoliation) by S. poliophaea caterpillars than their understorey counterparts. 4 Adding a novel dimension to the Janzen–Connell hypothesis, we found that early wet season herbivory of seedlings in gaps increased with conspecific adult density within a 125-m radius; whereas in the understorey only those seedlings within 50 m of a Swietenia tree were attacked by caterpillars. 5 Synthesis. These results suggest lepidopterans that need young leaves for food may forage more widely in forests to find seedlings in light-rich canopy gaps. Moths may achieve this successfully by being first attracted to gaps, and then searching within them for suitable hosts. A conceptual model, integrating conspecific adult tree density with light-driven changes in seedling tolerance/vigour and their susceptibility to herbivory and mortality, is presented. Spatial variation in the light available to tree seedlings often affects their tolerance and vigour, which may have important consequences for leaf-chewing insects and the scale of density-dependent herbivory in forests.

Remote detection and distinction of ants using nest-site specific LISS-derived Normalised Difference Vegetation Index

This study in Western Ghats, India, investigates the relation between nesting sites of ants and a single remotely sensed variable: the Normalised Difference Vegetation Index (NDVI). We carried out sampling in 60 plots each measuring 30 x 30 m and recorded nest sites of 13 ant species. We found that NDVI values at the nesting sites varied considerably between individual species and also between the six functional groups the ants belong to. The functional groups Cryptic Species, Tropical Climate Specialists and Specialist Predators were present in regions with high NDVI whereas Hot Climate Specialists and Opportunists were found in sites with low NDVI. As expected we found that low NDVI values were associated with scrub jungles and high NDVI values with evergreen forests. Interestingly, we found that Pachycondyla rufipes, an ant species found only in deciduous and evergreen forests, established nests only in sites with low NDVI (range = 0.015 - 0.1779). Our results show that these low NDVI values in deciduous and evergreen forests correspond to canopy gaps in otherwise closed deciduous and evergreen forests. Subsequent fieldwork confirmed the observed high prevalence of P. rufipes in these NDVI-constrained areas. We discuss the value of using NDVI for the remote detection and distinction of ant nest sites.

Spatial structuring of arbuscular mycorrhizal communities in benchmark and modified temperate eucalypt woodlands

Arbuscular mycorrhizal fungi (AMF) are crucial to the functioning of the plant–soil system, but little is known about the spatial structuring of AMF communities across landscapes modified by agriculture. AMF community composition was characterized across four sites in the highly cleared south-western Australian wheatbelt that were originally dominated by forb-rich eucalypt woodlands. Environmentally induced spatial structuring in AMF composition was examined at four scales: the regional scale associated with location, the site scale associated with past management (benchmark woodlands with no agricultural management history, livestock grazing, recent revegetation), the patch scale associated with trees and canopy gaps, and the fine scale associated with the herbaceous plant species beneath which soils were sourced. Field-collected soils were cultured in trap pots; then, AMF composition was determined by identifying spores and through ITS1 sequencing. Structuring was strongest at site scales, where composition was strongly related to prior management and associated changes in soil phosphorus. The two fields were dominated by the genera Funneliformis and Paraglomus, with little convergence back to woodland composition after revegetation. The two benchmark woodlands were characterized by Ambispora gerdemannii and taxa from Gigasporaceae. Their AMF communities were strongly structured at patch scales associated with trees and gaps, in turn most strongly related to soil N. By contrast, there were few patterns at fine scales related to different herbaceous plant species, or at regional scales associated with the 175 km distance between benchmark woodlands. Important areas for future investigation are to identify the circumstances in which recolonization by woodland AMF may be limited by fungal propagule availability, reduced plant diversity and/or altered chemistry in agricultural soils.

Effects of selective logging on tropical forest tree growth

We combined measurements of tree growth and carbon dioxide exchange to investigate the effects of selective logging on the Aboveground Live Biomass (AGLB) of a tropical rain forest in the Amazon. Most of the measurements began at least 10 months before logging and continued at least 36 months after logging. The logging removed similar to 15% of the trees with Diameter at Breast Height (DBH) greater than 35 cm, which resulted in an instantaneous 10% reduction in AGLB. Both wood production and mortality increased following logging, while Gross Primary Production (GPP) was unchanged. The ratio of wood production to GPP (the wood Carbon Use Efficiency or wood CUE) more than doubled following logging. Small trees (10 cm < DBH < 35 cm) accounted for most of the enhanced wood production. Medium trees (35 cm < DBH < 55 cm) that were within 30 m of canopy gaps created by the logging also showed increased growth. The patterns of enhanced growth are most consistent with logging-induced increases in light availability. The AGLB continued to decline over the study, as mortality outpaced wood production. Wood CUE and mortality remained elevated throughout the 3 years of postlogging measurements. The future trajectory of AGLB and the forest`s carbon balance are uncertain, and will depend on how long it takes for heterotrophic respiration, mortality, and CUE to return to prelogging levels.

Gap size measurement: the proposal of a new field method

In the tropical Atlantic Forest, 42 canopy gaps had their areas estimated using four different field methods of measurement: Runkle, Brokaw and Green [Runkle, J.R., 1981. Gap formation in some old-growth forests of the eastern United States. Ecology 62, 1041-1051; Brokaw, N.V.L., 1982. The definition of treefall gap and its effect on measures of forest dynamics. Biotropica 14, 158-160; Green, P.T., 1996. Canopy Gaps in rain forest on Christmas Island, Indian Ocean: size distribution and methods of measurement. J. Trop. Ecol. 12, 427-434] and a new method proposed in this work. It was found that within the same gap delimitation, average gap size varied from 56.0 up to 88.3 m(3) while total sum of gap area varied from 2351.3 to 3707.9 m(3) Differences among all methods and between pairs of method proved to be statistically significant. As a consequence, gap size-class distribution was also different between methods. When one method is held as a standard, deviation on average values of gap size ranged between 11.8 and 59.7% as deviations on single gap size can reach 172.8%. Implications on forest dynamics were expressed by the forest turnover rate that was 24% faster or 15% slower depending on the method adopted for gap measurement. Based on my results and on methods' evaluation, the use of a new method is proposed here for future research involving the measure of gap size in forest ecosystems. Finally, it is concluded that forest comparisons disregarding the influence of different methods of gap measurement should be reconsidered. (c) 2005 Elsevier B.V. All rights reserved.

Gap disturbance regime and composition in the Atlantic Montane Rain Forest: the influence of topography

In the Atlantic Montane Rain Forest of south-eastern Brazil, a field study was carried out to describe the forest disturbance regime, analyse canopy gap composition and evaluate the influence of habitat parameters on gap tree species composition. We characterized canopy gaps considering the group of variables as follows: area, type and number of tree/branch falls, topographic position, soil coverage and surrounding canopy trees. Gap composition was assessed at species level by measuring all individuals inside gaps higher than one meter. Mean gap area of the 42 canopy gaps analysed was 71.9 +/- 9.0 m(2) (mean +/- SE). Out of the studied gaps, 35.7% were created by uprooted and by snapped trees, 16.7% by dead-standing trees and 11.9% by the fall of large branches. The disturbance regime was characterized by gap openings predominantly smaller than 150 m(2) and by spatial patterning related to topography. Ridges had smaller gaps and higher proportions of gaps created by branch falls; slopes had bigger gaps generally created by uprooting events. The more abundant and frequent species were shade tolerant and the more species-rich families found inside gaps did not differ from the forest as a whole. Pioneer species were rare and restricted to medium and large size classes. The Indicator Species Analysis and the Canonical Correspondence Analysis indicated gap area, topography and the percentage of soil cover by the genera Calathea and Ctenanthe were the predominant variables correlated with woody species distribution. So, topography emerged as an important issue not only to the gap disturbance regime, but also to gap colonization. In respect to the influence of gap processes on the Atlantic Montane Rain Forest regeneration, our results support the view that canopy gap events may not be working as promoters of community wide floristic shifts.

Improving methods in gap ecology: Revisiting size and shape distributions using a model selection approach

Questions: We assess gap size and shape distributions, two important descriptors of the forest disturbance regime, by asking: which statistical model best describes gap size distribution; can simple geometric forms adequately describe gap shape; does gap size or shape vary with forest type, gap age or the method used for gap delimitation; and how similar are the studied forests and other tropical and temperate forests? Location: Southeastern Atlantic Forest, Brazil. Methods: Analysing over 150 gaps in two distinct forest types (seasonal and rain forests), a model selection framework was used to select appropriate probability distributions and functions to describe gap size and gap shape. The first was described using univariate probability distributions, whereas the latter was assessed based on the gap area-perimeter relationship. Comparisons of gap size and shape between sites, as well as size and age classes were then made based on the likelihood of models having different assumptions for the values of their parameters. Results: The log-normal distribution was the best descriptor of gap size distribution, independently of the forest type or gap delimitation method. Because gaps became more irregular as they increased in size, all geometric forms (triangle, rectangle and ellipse) were poor descriptors of gap shape. Only when small and large gaps (> 100 or 400m2 depending on the delimitation method) were treated separately did the rectangle and isosceles triangle become accurate predictors of gap shape. Ellipsoidal shapes were poor descriptors. At both sites, gaps were at least 50% longer than they were wide, a finding with important implications for gap microclimate (e.g. light entrance regime) and, consequently, for gap regeneration. Conclusions: In addition to more appropriate descriptions of gap size and shape, the model selection framework used here efficiently provided a means by which to compare the patterns of two different types of forest. With this framework we were able to recommend the log-normal parameters μ and σ for future comparisons of gap size distribution, and to propose possible mechanisms related to random rates of gap expansion and closure. We also showed that gap shape varied highly and that no single geometric form was able to predict the shape of all gaps, the ellipse in particular should no longer be used as a standard gap shape. © 2012 International Association for Vegetation Science.

Regeneração natural em ecounidades maduras e em reorganização de fragmentos de Floresta Estacional Semidecídua

The Atlantic Rainforest biome has been going through fragmentation processes caused by agriculture and urbanization in green areas. Structural studies associated with the silvigenetic approach allow the understanding of what the past has caused in the present structure and predict future conditions of disturbed fragments. The objective of this study was to compare the composition and diversity of arboreous natural regeneration of steady-state and reorganization ecounits in two Seasonal Semideciduous Forest fragments. The hypothesis was that specific composition varies in these two different ecounits due to differential adaptation of species in canopy gaps and closed canopy. The survey was made in three areas with different perturbation backgrounds of 0,5 ha each. 60 permanent plots of 4m² each (2m x 2m) were stablished along the studied fragments following the proportion of ecounits presented in a previous mapping. Each plot was divided in 4 sub-plots of 1m² and arboreous individuals between 0,20m and 1,30m height were sampled and posteriorly separated in two height classes: I) individuals between 0,20m and 0,50m height (2m² sampling) and II) individuals between 0,51m and 1,30m height (4m² sampling). It was sampled 338 individuals from 53 families and 23 species. The Shannon index was 3,26 (Area A), 2,27 (Area B) and 2,42 (Area C) whereas Areas B and C values are considered low in our state Semidecidous Forests. Steady-state ecounits presented the highest values for abundance and species richness. Chi-square test pointed out species’ selection for determined ecounits in the studied community. Rarefaction method analysis showed diversity increase in steady-state ecounits and a stablishment in species richness curves for reorganization ecounits

Spatial Relationships Among Young Cercocarpus ledifolius (Curlleaf Mountain Ylahogany)

This study analyzed spatial location patterns of Cercocarpus ledifolius Nutt. (curlleaf mountain mahogany) plants, classified as current-year seedling, established seedling, juvenile, and immature individuals, at a central Nevada study site. Most current-year seedlings were located in mahogany stands in which large, mature individuals had the greatest abundance. These stands had greater litter cover and a thicker layer of litter than areas with few current- year seedlings. Most established young Cercocarpus were located in adjacent Artemisia tridentata ssp. vaseyana (mountain big sagebrush) communities, or in infrequent canopy gaps between relatively few large, mature Cercocarpus. We discuss potential roles of plant litter, root growth characteristics, nurse plants, and herbivory in the establishment and renewal of Cercocarpus communities.

Seed fate and seedling dynamics after masting in two African rain forest trees

How the effects of biotic factors are moderated by abiotic factors, and their consequences for species interactions, is generally understudied in ecology. A key abiotic feature of forests is regular canopy disturbances that create temporary patches, or “gaps,” of above-average light availability. Co-occurring in lowland primary forest of Korup National Park (Cameroon), Microberlinia bisulcata and Tetraberlinia bifoliolata are locally dominant, ectomycorrhizal trees whose seeds share predator guilds in masting years. Here, we experimentally tested the impact of small mammal predators upon seedling abundance, growth, and survivorship. In 2007, we added a fixed density of seeds of each species to exclosures at 48 gap–understory locations across 82.5 ha within a large Microberlinia grove, and at 15 locations outside it. For both species, small mammals removed more seeds in gaps than in understory, whereas this was reversed for seeds killed by invertebrates. Nonetheless, Microberlinia lost twice as many seeds to small mammals, and more to invertebrates in exclosures, than Tetraberlinia, which was more prone to a pathogenic white fungus. After six weeks, both species had greater seedling establishment in gaps than understory, and in exclosures outside compared to exclosures inside the grove. In the subsequent two-year period, seedling growth and survivorship peaked in exclosures in gaps, but Microberlinia had more seedlings' stems clipped by animals than Tetraberlinia, and more than twice the percentage of leaf area damaged. Whereas Microberlinia seedling performance in gaps was inferior to Tetraberlinia inside the grove, outside it Microberlinia had reduced leaf damage, grew taller, and had many more leaves than Tetraberlinia. No evidence was found for “apparent mutualism” in the understory as seedling establishment of both species increased away from (>25 m) large stems of either species, pointing to “apparent competition” instead. In gaps, Microberlinia seedling establishment was lower near Tetraberlinia than conspecific adults because of context-dependent small mammal satiation. Stage-matrix analysis suggested that protecting Microberlinia from small mammals could increase its population growth rate by 0.06. In the light of prior research we conclude that small mammals and canopy gaps play an important role in promoting species coexistence in this forest, and that their strong interaction contributes to Microberlinia's currently very poor regeneration.

The influence of concurrent disturbances on plant community dynamics in northern hemlock-hardwood forests

Throughout the Upper Great Lakes region, alterations to historic disturbance regimes have influenced plant community dynamics in hemlock-hardwood forests. Several important mesic forest species, eastern hemlock (Tsuga canadensis), yellow birch (Betula alleghaniensis), eastern white pine (Pinus strobus), and Canada yew (Taxus canadensis), are in decline due to exploitive logging practices used at the turn of the 20th century and the wave of intense fires that followed. Continued regeneration and recruitment failure is attributed to contemporary forest management practices and overbrowsing by white-tailed deer (Odocoileus virginianus). Therefore, I examined the influence of two concurrent disturbances, overstory removal and herbivory, on plant community dynamics in two hemlock-hardwood forests. I measured the post-disturbance regeneration response (herbaceous and woody species) inside and outside of deer exclosures in 20 artificial canopy gaps (50 – 450 m2) and monitored survival and growth for hundreds of planted seedlings. The results of this research show that interacting disturbances can play a large role in shaping plant community composition and structure in hemlock-hardwood forests. White-tailed deer herbivory homogenized the post-disturbance plant communities across the experimental gradient of gap areas, essentially making species compositions in small gaps “look like” those in large gaps. Deer browsing also influenced probability of survival for planted Canada yew cuttings; all else being equal an individual was nearly seven times more likely to survive if protected from herbivory (P < 0.001). In contrast, the ability of sugar maple (Acer saccharum) to persist under high levels of herbivory and respond rapidly to overstory release appears to be related to the presence of stem layering(i.e., portions of below-ground prostrate stem). Layering occurred in 52% of excavated saplings (n = 100) and was significantly associated with increased post-disturbance height growth. Understory light was also important to planted seedling establishment and height growth. Higher levels of direct under-canopy light negatively impacted survival for shade-tolerant hemlock and Canada yew, while an increase in diffuse light was linked to a higher probability of survival for yellow birch and height growth for hemlock and Canada yew. Increases in white pine height growth were also significantly associated with a decrease in canopy cover.

Promoting Biodiversity in Forest Ecosystems Using Ecological Forestry

A subset of forest management techniques, termed ecological forestry, have been developed in order to produce timber and maintain the ecological integrity of forest communities through practices that more closely mirror natural disturbance regimes. Even though alternative methods have been described and tested, these approaches still need to be established and analyzed in a variety of geographic regions in order to calibrate and measure effectiveness across different forest types. The primary objective of this research project was to assess whether group selection combined with legacy-tree retention could enhance mid-tolerant tree recruitment in a late-successional northern hardwood forest. In order to evaluate a novel alternative regeneration technique, 49 group-selection openings in three size classes were created in 2003 with a biological legacy tree retained in the center of each opening. Twenty reference sites, managed using single-tree selection, were also analyzed for comparison. The specific goals of the project were to: 1) determine the fate and persistence of the openings and legacy trees 2) assess the understory response of the group-selection openings versus the single-tree selection reference sites, and 3) evaluate the spatial patterns of yellow birch (Betula alleghaniensis Britt.) and eastern hemlock (Tsuga canadensis (L.) Carr.) in the group-selection openings. The results from 8-9 years post-study implementation and the changes that have occurred between 2004/5 and 2011/12 are discussed. The alternative regeneration technique developed and assessed in this study has the potential to enrich biodiversity in a range of forest types. Projected group-selection opening persistence rates ranged from 41-91 years. Openings from 500-1500 m2 are predicted to persist long enough for mid-tolerant tree recruitment. The legacy trees responded well to release and experienced a low mortality rate. Yellow birch (the primary shade mid-tolerant tree in the study area) densities increased with opening size. Maples surpassed all other species in abundance. In the sapling layer, sugar maple (Acer saccharum Marsh.) was 2 to over 300 times more abundant in the group-selection openings and 2 to 3 times more abundant in the references sites than all other species present. Red maple (Acer rubrum L.) was the second most abundant species present in the openings and reference sites. Spatial patterns of yellow birch and eastern hemlock in the openings were mostly aggregated. The southern edges of the largest openings contained the highest magnitude of yellow birch and eastern hemlock per unit area. Continued monitoring and additional treatments will likely be necessary in order to ensure underrepresented species successfully reach maturity.