Effects of longleaf pine planting density and other factors on stand structure and associated wildlife habitat

The primary objective of many longleaf pine (Pinus palustris) restoration programs is to enhance or restore habitat for wildlife dependent on herbaceous plant communities. Because herbaceous cover is inversely related to canopy cover, restoration programs often place restrictions on longleaf pine planting density. However, the influence of planting density on understory plant communities has been inadequately evaluated. Therefore, we initiated a study to examine the relative influences of planting density and other factors on overall understory composition and forage availability for white‐tailed deer (Odocoileus virginianus) and northern bobwhite (Colinus virginianus) in nine longleaf pine stands throughout the Coastal Plain of Alabama during 2017–2018. We found that coverage of herbaceous plants decreased 3.5%, coverage of woody plants decreased 2.4%, and coverage of northern bobwhite forage plants decreased 1.9% for each 1 m²/ha increase in longleaf pine basal area. However, planting density was not a significant predictor of current basal area, nor coverage of any functional group of plants we examined, likely because current longleaf pine density averaged only 46% (range = 30–64%) of seedling planting density. We did not detect an effect of prescribed fire on stand condition or understory plant communities, likely due to variability in fire timing and frequency. Our findings related to planting density were likely a function of low longleaf pine survival, which is not uncommon. Because of this and the inherent variability in growth rates for young longleaf pine stands, restoration programs should consider placing greater emphasis on post‐planting monitoring and management than planting density.     

Hardwood management and restoration of longleaf pine ecosystems may affect raccoon daytime resting sites

Fire‐maintained Pinus palustris (longleaf pine) ecosystems are species rich and considered a top conservation priority in the southeastern United States. Ground‐nesting species such as Gopherus polyphemus (gopher tortoise) and Colinus virginianus (northern bobwhite) thrive in longleaf ecosystems. However, the generalist carnivore Procyon lotor (raccoon) is a significant predator of these endemic ground nesters. In forested ecosystems, raccoons prefer hardwood‐dominated habitats. Removal of hardwood trees, which is a common longleaf pine ecosystem restoration tool, affects habitat use of this predator. We examined 269 daytime resting sites (DRS) associated with 31 radio‐collared adult raccoons (18 M, 13 F) during 2014–2015 on a longleaf pine‐dominated site in southwestern Georgia. We developed 26 a priori models using an information theoretic approach to evaluate factors affecting use of DRS by raccoons. The top two models (ΔAIC < 2) had combined model weights of 0.75 and contained tree diameter, tree type, presence of nearby hardwood, and distances to pine, hardwood, mixed forest, and agriculture as predictors. However, the only informative variables were tree type and tree diameter. Raccoons used DRS in all available forest types, but were less likely to use pine trees (n = 7) relative to hardwoods (n = 247), and there was a positive relationship with tree diameter. Females used smaller trees farther from agriculture and primary roads, and were closer to wetlands than those used by males. Hardwood removal from within longleaf pine ecosystem affects habitat use of this predator, specifically DRS.     

Seed depredation negates the benefits of midstory hardwood removal on longleaf pine seedling establishment

Midstory hardwoods are traditionally removed to restore longleaf pine on fire‐excluded savannas. However, recent evidence demonstrating midstory hardwood facilitation on longleaf pine seedling survival has brought this practice into question on xeric sites. Also, midstory hardwoods could facilitate longleaf pine seedling establishment, as hardwood litter may conceal seeds from seed predators or improve micro‐environmental conditions for seedling establishment. However, little is known about these potential mechanisms. In this study, we tracked longleaf pine seed depredation and germination in artificially seeded plots (11 seeds/m²) in a factorial design fully crossing hardwood retention or removal with vertebrate seed predator access or exclusion in the Sandhills Ecoregion of North Carolina, U.S.A. Seed depredation averaged 78% across treatments and was greatest in unexcluded plots. Hardwood retention did not affect seed depredation. Longleaf pine averaged 3.6 germinants/4 m² across treatments, and was six times more abundant where vertebrates had been excluded. Hardwood removal had a strong positive effect on seedling germination, likely due to the removal of litter, but only when vertebrates were excluded. Our results indicated midstory hardwoods are not facilitating longleaf pine seedling establishment. Nevertheless, our results indicated that hardwood removal may not increase longleaf pine seedling establishment, as seed depredation diminished the effectiveness of hardwood removal under mast seed availability. Collectively, these results demonstrate the underlying complexity of the longleaf pine ecosystem, and suggest that planting may need to be part of the restoration strategy on sites where seed depredation limits longleaf pine natural regeneration.     

Distribution of native legumes (Leguminoseae) in frequently burned longleaf pine (Pinaceae)-wiregrass (Poaceae) ecosystems

Legume species distribution and abundance and selected environmental variables were quantified across a complex gradient (varying in both water-holding capacity and fertility) for frequently burned longleaf pine (Pinus palustris)–wiregrass (Aristida stricta) ecosystems. Legumes were present in all months; however, abundance peaked in June and was minimal after killing frosts in October. Legume species were prominent in the flora (43 species encountered) ubiquitous (94% of 2-m² subplots had at least one legume species), and abundant (nearly 120 000 stems/ha). Although most species were widely distributed throughout the gradient, Lespedeza angustifolia was distinctly associated with the more hydric end of the gradient, while both Petalostemon pinnatum and Galactia microphylla were located in the more xeric extreme. The percentage variation in species that could be accounted for by environmental variation was low (27%). Of the variation that could be accounted for, a number of environmental variables were important, including soil moisture, pine basal area (i.e., light), and bivalent base cations (e.g., Ca²⁺). Although gradients in resource availability among sites did not affect the distribution of species or abundance of legumes strongly, variation in resources are likely to regulate N₂-fixation rates of the various native legume species, and thereby affect ecological functions such as maintenance of N capital and productivity.     

Canopy thinning,not agricultural history,determines early responses of wild bees to longleaf pine savanna restoration

Longleaf pine savannas are highly threatened, fire‐maintained ecosystems unique to the southeastern United States. Fire suppression and conversion to agriculture have strongly affected this ecosystem, altering overstory canopies, understory plant communities, and animal populations. Tree thinning to reinstate open canopies can benefit understory plant diversity, but effects on animal communities are less well understood. Moreover, agricultural land‐use legacies can have long‐lasting impacts on plant communities, but their effects on animal communities either alone or through interactions with restoration are unclear. Resolving these impacts is important due to the conservation potential of fire‐suppressed and post‐agricultural longleaf savannas. We evaluated how historical agricultural land use and canopy thinning affect the diversity and abundance of wild bees in longleaf pine savannas. We employed a replicated, large‐scale factorial block experiment in South Carolina, where canopy thinning was applied to longleaf pine savannas that were either post‐agricultural or remnant (no agricultural history). Bees were sampled using elevated bee bowls. In the second growing season after restoration, thinned plots supported a greater bee abundance and bee community richness. Additionally, restored plots had altered wild bee community composition when compared to unthinned plots, indicating that reduction of canopy cover by the thinning treatment best predicted wild bee diversity and composition. Conversely, we found little evidence for differences between sites with or without historical agricultural land use. Some abundant Lasioglossum species were the most sensitive to habitat changes. Our results highlight how restoration practices that reduce canopy cover in fire‐suppressed savannas can have rapid benefits for wild bee communities.     

Influence of CO2 enrichment and nitrogen fertilization on tissue chemistry and carbon allocation in longleaf pine seedlings

One-year old, nursery-grown longleaf pine (Pinus palustris Mill.) seedlings were grown in 45-L pots containing a coarse sandy medium and were exposed to two concentrations of atmospheric CO₂ (365 or 720 mol^-1) and two levels of nitrogen (N) fertility (40 or 400 kg N ha^-1 yr^-1) within open top chambers for 20 months. At harvest, needles, stems, coarse roots, and fine roots were separated and weighed. Subsamples of each tissue were frozen in liquid N, lyophilized at -50°C, and ground to pass a 0.2 mm sieve. Tissue samples were analyzed for carbon (C), N, nonpolar extractives (fats, waxes, and oils = FWO), nonstructural carbohydrates (total sugars and starch), and structural carbohydrates (cellulose, lignin, and tannins). Increased dry weights of each tissue were observed under elevated CO₂ and with high N; however, main effects of CO₂ were significant only on belowground tissues. The high N fertility tended to result in increased partitioning of biomass aboveground, resulting in significantly lower root to shoot ratios. Elevated CO₂ did not affect biomass allocation among tissues. Both atmospheric CO₂ and N fertility tended to affect concentration of C compounds in belowground, more than aboveground, tissues. Elevated CO₂ resulted in lower concentrations of starch, cellulose, and lignin, but increased concentrations of FWO in root tissues. High N fertility increased the concentration of starch, cellulose, and tannins, but resulted in lower concentrations of lignin and FWO in roots. Differences between CO₂ concentrations tended to occur only with high N fertility. Atmospheric CO₂ did not affect allocation patterns for any compound; however the high N treatment tended to result in a lower percentage of sugars, cellulose, and lignin belowground.Joseph W. Jones     

Gopher Tortoise Response to Habitat Management by Prescribed Burning

ABSTRACT As quality of forested habitat declines from altered fire regimes, gopher tortoises (Gopherus polyphemus) often move into ruderal areas to the detriment of the animal and land manager. We evaluated effects of a dormant-and-growing-season prescribed fire on habitat and gopher tortoise use of degraded longleaf pine (Pinus palustris) forests surrounding military training areas. We burned 4 of 8 sites in winter 2001–2002 and again in April 2003. Changes in vegetation measured during 2001–2004 indicated that burn treatments did not increase herbaceous vegetation. Similarly, movement patterns, burrow usage, and home range of tortoises radiotracked from 2002–2004 did not differ between treatments. Woody cover initially was reduced in the forests postburn, and we found more new burrows in burned forest sites. Once shrub cover was reduced, tortoises started using forested habitat that had become overgrown. However, shrub reduction may be temporary, as woody stem densities increased postburn. Thus, the one-time use of fire to manage tortoise habitat may not rapidly restore the open canopy, sparse woody midstory, and abundant herbaceous vegetation that this species requires. Repeated prescribed fires or additional management techniques may be needed for complete restoration.     

Assessing the functional implications of soil biodiversity in ecosystems

Soil communities are among the most species-rich components of terrestrial ecosystems. A major challenge for soil ecologists is to formulate feasible research strategies that will preserve and capitalize on the biodiversity resources of the soil. This article considers the role of soil organism diversity by concentrating on: (i) the relationship between soil biodiversity and ecosystem function; (ii) what issues need to be explored; (iii) studies carried out in the Ecotron controlled environment facility; and (iv) how stable isotope techniques can improve our understanding of the relationship between soil biodiversity and ecosystem function. It is advocated that: (i) the objective of any soil biodiversity study should always be the generation of general concepts, rather than local, system-specific observations; and (ii) any empirical study can be properly interpreted only within a quantitative ecological framework.     

Ecological processes: A key element in strategies for nature conservation

Summary A common approach to nature conservation is to identify and protect natural ‘assets’ such as ecosystems and threatened species. While such actions are essential, protection of assets will not be effective unless the ecological processes that sustain them are maintained. Here, we consider the role of ecological processes and the complementary perspective for conservation arising from an emphasis on process. Many kinds of ecological processes sustain biodiversity: including climatic processes, primary productivity, hydrological processes, formation of biophysical habitats, interactions between species, movements of organisms and natural disturbance regimes. Anthropogenic threats to conservation exert their influence by modifying or disrupting these processes. Such threats extend across tenures, they frequently occur offsite, they commonly induce non‐linear responses, changes may be irreversible and the full consequences may not be experienced for lengthy periods. While many managers acknowledge these considerations in principle, there is much scope for greater recognition of ecological processes in nature conservation and greater emphasis on long time‐frames and large spatial scales in conservation planning. Practical measures that promote ecological processes include: monitoring to determine the trajectory and rate of processes; incorporating surrogates for processes in conservation and restoration projects; specific interventions to manipulate and restore processes; and planning for the ecological future before options are foreclosed. The long‐term conservation of biodiversity and the well‐being of human society depend upon both the protection of natural assets and maintaining the integrity of the ecological processes that sustain them.     

People as Ecological Participants in Ecological Restoration

In 1987, Bradshaw proposed that ecological restoration is the ultimate “acid test” of our understanding the functioning of ecosystems ( Bradshaw 1987 ). Although this concept is widely supported academically, how it can be applied by restoration practitioners is still unclear. This is an issue not limited to Bradshaw’s acid test, but moreover, reflects a general difficulty associated with the polarization between conceptual restoration (restoration ecology) and practical restoration (ecological restoration), where each has functioned to certain degree in isolation of the other. Outside of the more obvious pragmatic reasons for the relative independence between ecological restoration and restoration ecology, we propose that a more contentious explanation is that the approach taken toward understanding ecosystem development in restoration ecology is tangential to what actually takes place in ecological restoration. Current paradigms assume that the process of ecosystem development in restoration should follow the developmental trajectories suggested by classical ecological succession models. However, unlike these models, ecosystem development in restoration is, at least initially, largely manipulated by people, rather than by abiotic and biotic forces alone. There has been little research undertaken to explore how restoration activities impact upon or add to the extant ecological processes operating within a restoration site. Consequently, ecological restoration may not be so much an acid test of our understanding the functioning of ecosystems, but rather, an acid test of our understanding mutually beneficial interactions between humans and ecosystems.     

Atmospheric CO2 concentration, N availability, and water status affect patterns of ergastic substance deposition in longleaf pine (Pinus palustris Mill.) foliage

 Leaf chemistry alterations due to increasing atmospheric CO₂ will reflect plant physiological changes and impact ecosystem function. Longleaf pine was grown for 20 months at two levels of atmospheric CO₂ (720 and 365 μmol mol^–1), two levels of soil N (4 g m^–2 year^–1 and 40 g m^–2 year^–1), and two soil moisture levels (– 0.5 and – 1.5 MPa) in open top chambers. After 20 months of exposure, needles were collected and ergastic substances including starch grains and polyphenols were assessed using light microscopy, and calcium oxalate crystals were assessed using light microscopy, scanning electron microscopy, and transmission electron microscopy. Polyphenol content was also determined using the Folin-Denis assay and condensed tannins were estimated by precipitation with protein. Evaluation of phenolic content histochemically was compared to results obtained using the Folin-Denis assay. Total leaf polyphenol and condensed tannin content were increased by main effects of elevated CO₂, low soil N and well-watered conditions. Elevated CO₂ and low soil N decreased crystal deposition within needle phloem. Elevated CO₂ had no effect on the percentage of cells within the mesophyll, endodermis, or transfusion tissue which contained visible starch inclusions. With respect to starch accumulation in response to N stress, mesophyll > endodermis > transfusion tissue. The opposite was true in the case of starch accumulation in response to main effects of water stress: mesophyll < endodermis < transfusion tissue. These results indicate that N and water conditions significantly affect deposition of leaf ergastic substances in longleaf pine, and that normal variability in leaf tissue quality resulting from gradients in soil resources will be magnified under conditions of elevated CO₂. Received: 5 November 1996 / Accepted: 7 March 1997     

Carbon exchange of a mature,naturally regenerated pine forest in north Florida

We used eddy covariance and biomass measurements to quantify the carbon (C) dynamics of a naturally regenerated longleaf pine/slash pine flatwoods ecosystem in north Florida for 4 years, July 2000 to June 2002 and 2004 to 2005, to quantify how forest type, silvicultural intensity and environment influence stand‐level C balance. Precipitation over the study periods ranged from extreme drought (July 2000–June 2002) to above‐average precipitation (2004 and 2005). After photosynthetic photon flux density (PPFD), vapor pressure deficit (VPD) >1.5 kPa and air temperature <10 °C were important constraints on daytime half‐hourly net CO₂ exchange (NEE_day) and reduced the magnitude of midday CO₂ exchange by >5 μmol CO₂ m^?2 s^?1. Analysis of water use efficiency indicated that stomatal closure at VPD>1.5 kPa moderated transpiration similarly in both drought and wet years. Night‐time exchange (NEE_night) was an exponential function of air temperature, with rates further modulated by soil moisture. Estimated annual net ecosystem production (NEP) was remarkably consistent among the four measurement years (range: 158–192 g C m^?2 yr^?1). In comparison, annual ecosystem C assimilation estimates from biomass measurements between 2000 and 2002 ranged from 77 to 136 g C m^?2 yr^?1. Understory fluxes accounted for approximately 25–35% of above‐canopy NEE over 24‐h periods, and 85% and 27% of whole‐ecosystem fluxes during night and midday (11:00–15:00 hours) periods, respectively. Concurrent measurements of a nearby intensively managed slash pine plantation showed that annual NEP was three to four times greater than that of the Austin Cary Memorial Forest, highlighting the importance of silviculture and management in regulating stand‐level C budgets.     

生态能质(eco-exergy)在水生生态系统建模和评价中的应用

生态能质(eco-exergy)是指系统从给定状态达到热力学平衡状态所做的功,可以度量生态系统的复杂生物化学组分及生态结构。系统具有的能质越大,有序化程度越高,稳定性也越强。生态能质和比生态能质(specific eco-exergy)指标能够评价水生生态系统健康状况,对水生生态系统演替阶段具有指示作用。本文阐述了生态能质的定义、生态学意义以及生态能质值和权重因子fi值的计算方法。对生态能质作为功能函数在水生生态系统结构动力学模型(structural dynamic models,SDMs)中以及作为生态指标在生态系统健康评价、生态恢复研究中的实际应用进行了总结。最后探讨了生态能质指标的局限性以及未来的发展。此外,文中建议将eco-exergy和specific eco-exergy统一翻译为生态能质和比生态能质。     

Ecological Restoration Treatments Increase Butterfly Richness and Abundance: Mechanisms of Response

Few ecosystem restoration studies evaluate whether arthropods are important components of ecosystem recovery. We tested the hypothesis that ponderosa pine restoration treatments would increase adult butterfly species richness and abundance as a direct result of increased understory diversity and abundance. To examine mechanisms that potentially affect adult butterfly distribution, we quantified host plant frequency, nectar plant abundance, and insolation (light intensity) in restoration treatment and control forests. This study is unique, because this is the first invertebrate monitoring in ponderosa pine forest restoration treatments in the U.S. Southwest and also because these treatments are the first replicated ponderosa pine restoration treatments at a landscape scale. Three patterns emerged: (1) butterfly species richness and abundance were 2 and 3 times greater, respectively, in restoration treatment units than in paired control forests 1 year after treatment, and 1.5 and 3.5 times greater, respectively, 2 years after treatment, ordination of control and treatment sampling units using butterfly assemblages showed significant separation of control and restoration treatment units after restoration treatment; (2) host plant and nectar plant species richness showed little difference between treated and control forests even 2 years after treatment; and (3) insolation (light intensity) was significantly greater in treated forests after restoration. We suggest that changes in the butterfly assemblage may occur due to light intensity effects before plant community changes occur or can be detected. Butterfly assemblage differences will have additional cascading effects on the ecosystem as prey for higher trophic levels and through plant interactions including herbivory and pollination.     

Ecological consequences of altered hydrological regimes in fragmented ecosystems in southern Australia: Impacts and possible management responses

Abstract Although the potential impacts of rising water tables and secondary salinization on agricultural land in southern Australia have been recognized for some time, it is only recently that the impacts on native vegetation have been considered. Despite the likely extent and severity of the problem, no comprehensive approach to assessing the impact of salinity upon native vegetation has been attempted to date. In the present paper, we discuss the causes and impacts of rising water tables and dryland salinity, assess the levels of risk in different ecosystem types and consider the possibilities for the maintenance of biodiversity and ecosystem function in vegetation at risk. We examine the salinity risk to woodland vegetation in the Western Australian (WA) wheatbelt, and consider both broad‐scale context and finer‐scale variation within individual patches of vegetation. From this information, we develop a set of conceptual models of the potential impacts of shallow saline water tables on ecosystem structure and processes in remnant vegetation in agricultural areas, particularly in the WA wheatbelt. First, we suggest that fine‐scale variability in surface topography and soil characteristics may play an important role in limiting the impacts of rising saline water tables. The outcome will depend on the interaction of the heterogeneity of the impact, species distribution in relation to small‐scale environmental heterogeneity and variation in species response to hydrological change. Second, we suggest that shallow saline water tables can be considered to cause an ‘edge effect’, which moves inwards from the edge of remnants of native vegetation. Finally, we consider how saline surface flows exacerbate the effects of shallow saline water tables and hasten vegetation decline in remnant areas. We put these models forward as hypotheses to be tested in different situations. We contrast the situation of secondary salinization in Australian vegetation with that of naturally saline systems in Australia and elsewhere, and suggest that these systems may provide important signposts toward developing management approaches for vegetation at risk. In conclusion, we consider the need to set priorities for the protection and restoration of natural vegetation at risk from altered hydrology, based on an assessment of relative threat and probability of persistence or recovery. We highlight the urgency for action that protects native vegetation from the increasing risks of rising water tables.     

不同生态恢复方式下生态系统服务与生物多样性恢复效果的整合分析

全球范围内关键生态系统服务的减少使人类社会面临巨大的威胁,生物多样性是生态系统提供各种产品和服务的基础。生态恢复工程对退化的生态系统服务和生物多样性进行修复,对于缓解人类环境压力具有非常重要的意义。长期的理论和实践工作形成了多种生态恢复措施:(1)单纯基于生态系统自我设计的自然恢复方式,(2)人为设计对环境条件进行干预,反馈影响生态系统的自我设计,(3)人为设计对目标种群和生态系统进行直接干预和重建。这3类恢复方式可以在不同程度上定向的影响生态系统的恢复进程,反映了人类对生态系统的低度、中度和高度介入。哪种恢复方式和介入程度能够实现更好的恢复效果,是生态恢复学中的一个关键问题,但到目前为止,虽广有争议,却无定量的分析和结论。针对这个空白,通过对ISI Web of Knowledge数据库中生态恢复相关文献的整合分析,基于数学统计的方法定量比较在不同条件下低度介入(自然恢复)、中度介入(环境干预)和高度介入(直接干预)3种恢复方式对生态系统服务与生物多样性的恢复效果。论文从4个方面展开研究:(1)低度、中度、高度介入生态恢复方式的划分,(2)比较3大类介入方式对生态系统服务和生物多样性恢复效果的差异,(3)不同气候条件、生态系统类型和恢复时间等背景因素的影响,(4)生物多样性恢复和生态系统服务恢复之间的关系。研究结果揭示了不同生态恢复方式的适用条件,以及对生物多样性和生态系统恢复相互关系的作用,对生态恢复实践中恢复方式的选择有指导作用。对未来的研究也有启示意义,如针对特定生态系统服务或具体研究问题进一步探索低度、中度和高度介入生态恢复方式的作用规律和机制;将地区的社会经济水平、生态系统的受损程度等因素纳入生态恢复方式的考察,以最优化生态恢复成本-效率等。