Understory vegetation response to mountain pine beetle disturbance in northern Colorado lodgepole pine forests

Understory plants are an important element of forests, having a considerable influence on ecosystem functioning and canopy-tree development following disturbance. Recent bark beetle outbreaks across western North American forests have caused extensive canopy mortality, creating new growing conditions that provide the opportunity for changes within the intact understory. Over a five-year period following peak mountain pine beetle (MPB) activity across lodgepole pine-dominated forests in Rocky Mountain National Park, Colorado, we measured the changes in plant diversity, cover, and dominance by lifeform and quantified tree regeneration rates. Average species richness and diversity increased, but overall plant cover did not change. Graminoids appeared to benefit the most, increasing in average cover, richness, and relative dominance. The rise in graminoid dominance was largely at the expense of shrubs, which showed little ability to benefit from overstory mortality within the first years following attack. Most plant responses were positively related to the total tree basal area lost since the peak of the outbreak, suggesting that increased resource availability following tree death may benefit understory plants. However, a negative relationship between several understory variables and tree sapling density provides evidence that understory plants compete with saplings for the newly available resources. Tree seedling density nearly doubled over the duration of the study, indicating a strong regeneration pulse. Among species, lodgepole pine displayed the greatest tree seedling establishment. This is one of the first studies to use repeated measurements to describe this often-overlooked component of forest change associated with MPB disturbance.

     

The response of herbaceous-layer vegetation to anthropogenic disturbance in intermittent stream bottomland forests of southern Indiana,USA

Between 1993 and 1995 we sampled herbaceous layer vegetation on 84 plots in Platanus/Asarum Wet-Mesic Bottomland forests to determine how these forests have responded to human disturbance. Four different disturbance types were sampled (abandoned agricultural are as, clearcuts, group-selection openings, and single-tree selection openings), and uncut 80–100 year-old reference stands were sampled for comparison. Detrended Correspondence Analysis (DCA), distance analyses (chord distance and normalized Euclidean distance) and similarity analysis (Bray-Curtis similarity coefficient) suggest that agricultural use has shifted herbaceous-layer vegetation composition away from that typical of the reference forests, but that clearcutting, group-selection harvest, and single-tree selection harvest have not greatly shifted herbaceous composition. This shift in vegetation on abandoned agricultural land resulted from a loss of indicator species, such as Cardamine concatenata (Michx.) Sw., Stellaria pubera Michx., and Laportea canadensis (L.) Weddell and an influx of disturbance, exotic, and nonforest species (e.g., Lycopodium complanatum L., Lonicera japonica Thunb. and Senecio aureus L.). However, only two species found in reference stands, Erigenia bulbosa (Michx.) Nutt. and Sphenopholis obtusata (Michx.) Scribn., were missing from clearcuts, group-selection openings, and single-tree selection openings. The species richness values of abandoned agriculture, clearcut, and group-selection plots were generally greater than those of single-tree selection and reference plots. Abandoned agricultural areas had much greater total species richness because of the influx of dry-site, exotic, disturbance, and non-forest species.     

Territoriality of Feral Pigs in a Highly Persecuted Population on Fort Benning,Georgia

ABSTRACT We examined home range behavior of female feral pigs (Sus scrofa) in a heavily hunted population on Fort Benning Military Reservation in west-central Georgia, USA. We used Global Positioning System location data from 24 individuals representing 18 sounders (i.e., F social groups) combined with mark-recapture and camera-trap data to evaluate evidence of territorial behavior at the individual and sounder levels. Through a manipulative experiment, we examined evidence for an inverse relationship between population density and home range size that would be expected for territorial animals. Pigs from the same sounder had extensive home range overlap and did not have exclusive core areas. Sounders had nearly exclusive home ranges and had completely exclusive core areas, suggesting that female feral pigs on Fort Benning were territorial at the sounder level but not at the individual level. Lethal removal maintained stable densities of pigs in our treatment area, whereas density increased in our control area; territory size in the 2 areas was weakly and inversely related to density of pigs. Territorial behavior in feral pigs could influence population density by limiting access to reproductive space. Removal strategies that 1) match distribution of removal efforts to distribution of territories, 2) remove entire sounders instead of individuals, and 3) focus efforts where high-quality food resources strongly influence territorial behaviors may be best for long-term control of feral pigs.     

Soil properties as indicators of disturbance in forest ecosystems of Georgia,USA

The objective of this study was to examine spatial variability and the effects of forest ecosystem disturbance, caused by military training, on soil organic matter (SOM) and related soil properties. Six 5th to 6th order watersheds were sampled at the Fort Benning military reservation, Georgia, USA. Sampling locations were categorized as either upland or bottomland, and sites were classified as low, moderately or severely impacted according to the level of ecosystem disturbance. Total carbon (C) and nitrogen (N), and extractable iron (Fe) and aluminum (Al) were greater in bottomland sites. Extractable-cations were affected by disturbance on upland soil. Severely impacted sites showed lower total C and N concentrations in both landscape positions. This is likely due to the physical disruption of soil structure due to the traffic of heavy military vehicles, resulting in the removal of forest floor. This pattern was more evident in upland sites. Conversely, soil properties in the bottomland sites were less affected by the level of impact.     

Detecting anthropogenic disturbance in tropical forests

Tropical forests are beleaguered by an array of threats driven by different scales of anthropogenic perturbations, which vary in the degree to which they can be detected by remote sensing. The extent of different patterns of cryptic disturbance often far exceeds the total area deforested, as shown by two recent studies on selective logging in Amazonia. Here, we discuss different forms of disturbance in Amazonian forests and question how much of the apparently intact forest in this region remains relatively undisturbed.     

Selecting indicators of soil, microbial, and plant conditions to understand ecological changes in Georgia pine forests

Characterizing how resource use and management activities affect ecological conditions is necessary to document and understand anthropogenic changes in ecological systems. Resource managers on military installations have the delicate task of balancing the training needs of soldiers effectively with the need to maintain a high quality of ecological conditions. This study considers ways that ecological indicators can provide information on impacts that training has on environmental characteristics that occur at different scales and in different sectors of the environment. The characteristics examined include soil chemistry, soil microbes, and vegetation. A discriminant function analysis was conducted to determine whether ecological indicators could differentiate among different levels of military use. A combination of 10 indicators explained 90% of the variation among plots from five different military use levels. Results indicated that an appropriate suite of ecological indicators for military resource managers includes soil, microbial, and vegetation characteristics. Since many of these indicators are related, managers at this location potentially have freedom to choose indicators that are relatively easy to measure, without sacrificing information.     

Changes in Soil Properties Following 55 Years of Secondary Forest Succession at Fort Benning,Georgia, U.S.A.

We present results on changes in soil properties following land use change over an approximately 55‐year period at Fort Benning, Georgia, U.S.A. Soil cores were taken at 129 locations that were categorized as reforested (field/bare ground in 1944 and forest in 1999), disturbed (field/bare ground in 1944 and 1999), or reference forests (forest in 1944 and 1999). Soil disturbance included historic agriculture (pre‐1944) and military training (post‐1944). Density in mineral soils exhibited a historic land use legacy effect (reference < reforested < disturbed). Rates of change in bulk density decreased with depth and estimated total times to reach reference forest levels ranged from 83 (0–10 cm) to 165 (30–40 cm) years. A land use legacy effect on C stock was apparent in the O‐horizon and in 30‐ to 40‐cm soil increment (reference > reforested > disturbed). Soil C stock in all other increments and in particulate organic matter was affected by disturbance; however, no legacy was apparent (reference = reforested > disturbed). For the entire soil profile (O‐horizon to 40 cm), rate of C accrual was 28 g m⁻² yr⁻¹ (1.5%/yr). Nitrogen stocks were affected by disturbance in the O‐horizon and 0‐ to 10‐cm increment; however, no legacy effect was detected (reference = reforested > disturbed). Nitrogen accumulated at 0.56 g m⁻² yr⁻¹ (0.6%/yr) for the entire soil profile. At Fort Benning, soil C and N stocks of reforested stands were similar to those of reference forested stands after approximately 55 years. However, soil bulk density was greater on reforested stands than reference forest stands at 55 years and may require an additional century to reach reference levels.     

Effects of fire season on flowering of forbs and shrubs in longleaf pine forests

Summary Effects of variation in fire season on flowering of forbs and shrubs were studied experimentally in two longleaf pine forest habitats in northern Florida, USA. Large, replicated plots were burned at different times of the year, and flowering on each plot was measured over the twelve months following fire. While fire season had little effect on the number of species flowering during the year following fire, fires during the growing season decreased average flowering duration per species and increased synchronization of peak flowering times within species relative to fires between growing seasons. Fires during the growing season also increased the dominance of fall flowering forbs and delayed peak fall flowering. Differences in flowering resulting from variation in fire season were related to seasonal changes in the morphology of clonal forbs, especially fall-flowering composites. Community level differences in flowering phenologies indicated that timing of fire relative to environmental cues that induced flowering was important in determining flowering synchrony among species within the ground cover of longleaf pine forests. Differences in fire season produced qualitatively similar effects on flowering phenologies in both habitats, indicating plant responses to variation in the timing of fires were not habitat specific.     

Understory vegetation response to thinning disturbance of varying complexity in coniferous stands

Question: Can augmented forest stand complexity increase understory vegetation richness and cover and accelerate the development of late‐successional features? Does within‐stand understory vegetation variability increase after imposing treatments that increase stand structural complexity of the overstory? What is the relative contribution of individual stand structural components (i.e. forest matrix, gaps, and leave island reserves) to changes in understory vegetation richness? Location: Seven study sites in the Coastal Range and Cascades regions of Oregon, USA. Methods: We examined the effects of thinning six years after harvest on understory plant vascular richness and cover in 40‐ to 60‐year‐old forest stands dominated by Douglas‐fir (Pseudotsuga menziesii). At each site, one unthinned control was preserved and three thinning treatments were implemented: low complexity (LC, 300 trees ha^?1), moderate complexity (MC, 200 trees ha^?1), and high complexity (HC, variable densities from 100 to 300 trees ha^?1). Gaps openings and leave island reserves were established in MC and HC. Results: Richness of all herbs, forest herbs, early seral herbs and shrubs, and introduced species increased in all thinning treatments, although early seral herbs and introduced species remained a small component. Only cover of early seral herbs and shrubs increased in all thinning treatments whereas forest shrub cover increased in MC and HC. In the understory, we found 284 vascular plant species. After accounting for site‐level differences, the richness of understory communities in thinned stands differed from those in control stands. Within‐treatment variability of herb and shrub richness was reduced by thinning. Matrix areas and gap openings in thinned treatments appeared to contribute to the recruitment of early seral herbs and shrubs. Conclusions: Understory vegetation richness increased 6 years after imposing treatments, with increasing stand complexity mainly because of the recruitment of early seral and forest herbs, and both low and tall shrubs. Changes in stand density did not likely lead to competitive species exclusion. The abundance of potentially invasive introduced species was much lower compared to other plant groups. Post‐thinning reductions in within‐treatment variability was caused by greater abundance of early seral herbs and shrubs in thinned stands compared with the control. Gaps and low‐density forest matrix areas created as part of spatially variably thinning had greater overall species richness. Increased overstory variability encouraged development of multiple layers of understory vegetation.     

Mountain pine beetle selectivity in old‐growth ponderosa pine forests,Montana, USA

A historically unprecedented mountain pine beetle (MPB) outbreak affected western Montana during the past decade. We examined radial growth rates (AD 1860–2007/8) of co‐occurring mature healthy and MPB‐infected ponderosa pine trees collected at two sites (Cabin Gulch and Kitchen Gulch) in western Montana and: (1) compared basal area increment (BAI) values within populations and between sites; (2) used carbon isotope analysis to calculate intrinsic water‐use efficiency (iWUE) at Cabin Gulch; and (3) compared climate‐growth responses using a suite of monthly climatic variables. BAI values within populations and between sites were similar until the last 20–30 years, at which point the visually healthy populations had consistently higher BAI values (22–34%) than the MPB‐infected trees. These results suggest that growth rates two–three decades prior to the current outbreak diverged between our selected populations, with the slower‐growing trees being more vulnerable to beetle infestation. Both samples from Cabin Gulch experienced upward trends in iWUE, with significant regime shifts toward higher iWUE beginning in 1955–59 for the visually healthy trees and 1960–64 for the MPB‐infected trees. Drought tolerance also varied between the two populations with the visually healthy trees having higher growth rates than MPB‐infected trees prior to infection during a multi‐decadal period of drying summertime conditions. Intrinsic water‐use efficiency significantly increased for both populations during the past 150 years, but there were no significant differences between the visually healthy and MPB‐infected chronologies.     

Effects of mountain pine beetle on fuels and expected fire behavior in lodgepole pine forests, Colorado, USA

In Colorado and southern Wyoming, mountain pine beetle (MPB) has affected over 1.6 million ha of predominantly lodgepole pine forests, raising concerns about effects of MPB-caused mortality on subsequent wildfire risk and behavior. Using empirical data we modeled potential fire behavior across a gradient of wind speeds and moisture scenarios in Green stands compared three stages since MPB attack (Red [1-3 yrs], Grey [4-10 yrs], and Old-MPB [～30 yrs]). MPB killed 50% of the trees and 70% of the basal area in Red and Grey stages. Across moisture scenarios, canopy fuel moisture was one-third lower in Red and Grey stages compared to the Green stage, making active crown fire possible at lower wind speeds and less extreme moisture conditions. More-open canopies and high loads of large surface fuels due to treefall in Grey and Old-MPB stages significantly increased surface fireline intensities, facilitating active crown fire at lower wind speeds (>30-55 km/hr) across all moisture scenarios. Not accounting for low foliar moistures in Red and Grey stages, and large surface fuels in Grey and Old-MPB stages, underestimates the occurrence of active crown fire. Under extreme burning conditions, minimum wind speeds for active crown fire were 25-35 km/hr lower for Red, Grey and Old-MPB stands compared to Green. However, if transition to crown fire occurs (outside the stand, or within the stand via ladder fuels or wind gusts >65 km/hr), active crown fire would be sustained at similar wind speeds, suggesting observed fire behavior may not be qualitatively different among MPB stages under extreme burning conditions. Overall, the risk (probability) of active crown fire appears elevated in MPB-affected stands, but the predominant fire hazard (crown fire) is similar across MPB stages and is characteristic of lodgepole pine forests where extremely dry, gusty weather conditions are key factors in determining fire behavior.     

Characterization of Native American vegetation disturbance in the forests of central New York State,USA during the late 18th century ce

Vegetation History and Archaeobotany - Historic land survey records (LSRs) offer important details on local- and landscape-scale vegetation patterns related to Native American land-use practices...     

Alien plant species composition and associations with anthropogenic disturbance in North American forests

A probability-based sampling scheme was used to survey plant species composition in forests of 16 states in seven geopolitical regions of the United States (California, Colorado, Minnesota, and parts of the Pacific Northwest, Southeast, Mid-Atlantic, and Northeast) in 1994. The proportion of alien species relative to the total species number and to canopy cover in the ground stratum (0–0.6 m height) was evaluated in 279 plots. Visually evident anthropogenic disturbances (e.g., artificial regeneration, logging, prescribed burning, and grazing by livestock), if any, were recorded on each plot. In each of the seven regions we quantified (1) the percentage of the number of species and total cover comprised of alien species, (2) the difference in these percentages for disturbed and undisturbed plots, and (3) the origin or native range for the alien species.The percentage of alien species ranged from approximately 4.5% (Colorado) to approximately 13.2% (California). The percentage of alien species cover ranged from approximately 1.5% in Colorado to 25% in California. In five regions, species introduced from temperate Eurasia comprised the largest proportion of alien species and cover. In the Southeast, species introduced from far eastern and subtropical Asia dominated the alien flora. In the Mid-Atlantic, the majority of alien species was Eurasian and the majority of alien species cover consisted of far eastern and subtropical Asian species.The proportion of plots in which at least one alien species was recorded was significantly higher in disturbed than undisturbed plots in the Southeast and marginally significantly higher ($p=0.053$) in the Northeast. These results are consistent with other published studies that indicate that anthropogenic disturbance affects the structure and composition of both the ground stratum and upper canopy of forest habitats. In other regions, however, no significant differences were found.     

Understory vegetation response to mechanical mastication and other fuels treatments in a ponderosa pine forest

Questions: What influence does mechanical mastication and other fuel treatments have on: (1) canopy and forest floor response variables that influence understory plant development; (2) initial understory vegetation cover, diversity, and composition; and (3) shrub and non‐native species density in a second‐growth ponderosa pine forest. Location: Challenge Experimental Forest, northern Sierra Nevada, California, USA. Methods: We compared the effects of mastication only, mastication with supplemental treatments (tilling and prescribed fire), hand removal, and a control on initial understory vegetation response using a randomized complete block experimental design. Each block (n=4) contained all five treatments and understory vegetation was surveyed within 0.04‐ha plots for each treatment. Results: While mastication alone and hand removal dramatically reduced the midstory vegetation, these treatments had little effect on understory richness compared with control. Prescribed fire after mastication increased native species richness by 150% (+6.0 species m²) compared with control. However, this also increased non‐native species richness (+0.8 species m²) and shrub seedling density (+24.7 stems m²). Mastication followed by tilling resulted in increased non‐native forb density (+0.7 stems m²). Conclusions: Mechanical mastication and hand removal treatments aided in reducing midstory fuels but did not increase understory plant diversity. The subsequent treatment of prescribed burning not only further reduced fire hazard, but also exposed mineral soil, which likely promoted native plant diversity. Some potential drawbacks to this treatment include an increase of non‐native species and stimulation of shrub seed germination, which could alter ecosystem functions and compromise fire hazard reduction in the long‐term.     

Using species abundance models as indicators of habitat disturbance in tropical forests

1. In temperate communities, species abundance distributions have been used to detect ecosystem disturbance: in undisturbed habitats, distributions are claimed to generally fit log-normal models, whereas in disturbed habitats, distributions fit log-series models.
2. There is a growing literature on the effects of habitat disturbance in tropical ecosystems and several studies suggest that species abundance models may be useful in detecting disturbance, although data are lacking.
3. Nummelin (1998) claims that these models are not universal indicators of forest disturbance, but we highlight a number of problems with the data presented in Nummelin's study and conclude that it is too soon to dismiss these models. We discuss several important points arising from Nummelin's study which need to be considered if species abundance models are to be used appropriately.     

Habitat indicators for cavity-nesters: The polypore Phellinus pini in pine forests

Foresters and arborists have long used fruit-bodies of heart-rot fungi as signs of advanced live-tree decay, but such usage has not been elaborated for forest conservation. I analysed relationships between a heart-rot fungus, tree-cavity supply, and cavity-nesting bird assemblage in wet hemiboreal Scots pine (Pinus sylvestris) forests in Estonia. The focal species, Phellinus pini, is the main heartwood decayer of live pines; it typically forms fruit-bodies at the stage of advanced decay on old trees. I found that the pine wetlands had few tree-cavities (mostly in snags) and cavity-nesting birds (woodpeckers being almost absent) despite abundant snag supply. Only one fruit-body of P. pini was found on cavity-tree but stand-scale abundance of the fruit-bodies correlated well with cavity-nester densities. Multifactor models indicated that cavity formation, not tree death, was the limiting process for secondary cavity-nesters, and P. pini could indeed be used (in combination with old-pine abundance) for assessing their habitat quality. This fungus could also serve as an educational flagship species to bridge conservation biology and forest pathology, and its fruit-bodies can signal trees to be retained at harvesting in pine forests. The conclusion is that there is hope for developing practical indicators to manage for the hidden decay processes that govern tree-cavity development.     

Understory development in canopy gaps of pine and pine-hardwood forests of the upper Coastal Plain of Virginia

Canopygaps are important in establishing a pool of natural regeneration in manytemperate forest ecosystems. Information on the role of gaps in loblolly pine(Pinus taeda L.) and pine-hardwood foreststands in the southeastern Coastal Plain of the United States is lacking.Accordingly, 12 small canopy gaps in mature pine and pine-hardwood standsin Petersburg National Battlefield, Virginia, were studied. Loblolly pineregeneration was significantly more abundant in canopy gaps as compared to theadjacent forest in both forest cover types. In four stands dominated by loblollypine, there were 750 saplings/ha in the gaps compared to only 125saplings/ha in the adjacent forest. Pine saplings dominated the regenerationspectrum in the gaps in the pine stands, while red maple (Acerrubrum L.) was more important in the adjacent forest. In fourpine-hardwood stands, regeneration in both the gaps and adjacent forestwas dominated by sweetgum (Liquidambar styracifluaL.) with importance values of 27% and 28%, respectively.There were no loblolly pine seedlings in the adjacent forest, but an average 313per ha in the gaps of the pine-hardwood stands. Within thegaps in both cover types, loblolly pine saplings were lower in stature thancompeting hardwood stems, leading to the conclusion that the gaps may form atemporary pool of pine regeneration. Without further stand disturbance, theprocess of gap closure may reduce the pine component to a secondary status, orperhaps eliminate it altogether.     

Effects of anthropogenic disturbance on the species assemblages of birds in the back mangrove forests

Wetlands Ecology and Management - Around the world, mangrove ecosystem has faced major threats from human activities, and birds were unexceptionally impacted. Mangrove fringe a large extent of the...