Effects of habitat and growing season fires on resprouting of shrubs in longleaf pine savannas

The effects of habitat and timing of growing season fires on resprouting of shrubs were studied in second-growth longleaf pine savannas of the west Gulf coastal plain in the southeastern United States. Within the headwaters of three different drainages of the Calcasieu River in the Kisatchie National Forest in western Louisiana, replicated permanent transects were established that extended from xeric upland longleaf pine savannas into downslope hydric seepage savannas. All shrubs were mapped and tagged, and numbers of stems were counted prior to any fires. Replicated prescribed fires were set early (June) and late (August) during the 1990 growing season; maximum fire temperatures were measured within both upland and seepage habitats within each transect. Shrubs were relocated; stems were recensused two and twelve months after the fires. At least some shrubs of all species resprouted from underground organs; none regenerated solely from seed banks in the soil. There was no reduction in total numbers of stems one year after fires compared to before fires, either in the upland or in seepage savannas. In addition, there was no reduction in total numbers of stems one year after early or late growing season fires. Fire-related mortality was restricted to small shrubs (< 18 stems) and was not associated with high fire temperatures. The rate of resprouting varied among species and between habitats. Resprouting occurred more rapidly in seepage than upland savannas, but more resprouts were produced in upland than seepage savannas one year after fires. In contrast to other upland species, Vaccinium arboreum and V. elliottii delayed resprouting more than two months following fire. Stems of Rhus copallina and Pyrus arbutifolia, species with long rhizomes, increased more after fires in June than fires in August. We suggest that growing season fires may block further recruitment of shrubs into longleaf pine savannas, but reduction in numbers of large shrubs may require additional management.     

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.     

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.     

Fire in the Suburbs: Ecological Impacts of Prescribed Fire in Small Remnants of Longleaf Pine (Pinus palustris) Sandhill

Abstract Logging, fire suppression, and urbanization have all contributed to the serious decline and fragmentation of Pinus palustris (longleaf pine) ecosystems in the southeastern United States. Effective management of the remaining patches of these pyrogenic communities must incorporate periodic low‐intensity fires, even where they are located on private lands in populated urban and suburban areas. To explore the effects of fire and its potential use for restoration and management of small fragments surrounded by suburban development, we conducted growing season prescribed fires in remnant longleaf pine sandhill patches in the suburbs of Gainesville, Florida. Density and composition of hardwoods were surveyed pre‐burn and 1 and 9 months post‐burn. Woody stem density decreased in the burn plots, predominantly in the smaller size classes. Flowering responses of forbs and small shrubs were surveyed six times post‐burn for 1 year. Overall, the burns did not yield greater densities of flowering stems, but burn patches had higher species richness and diversity than control patches. In addition, there were consistently greater numbers of “showy flowered” sandhill species in flower in burn patches relative to controls. The results of this research demonstrate that prescribed fire can be used for restoration and management of small remnants of longleaf pine sandhill in suburban neighborhoods. It is also clear that although a single prescribed burn can be effective, it will take more than one burn to attain desired restoration goals in degraded longleaf remnants.     

Burning Influences on Wiregrass (Aristida beyrichiana) Restoration Plantings: Natural Seedling Recruitment and Survival

Regeneration and expansion of Aristida beyrichiana and Aristida stricta (wiregrass) populations in remaining fire‐maintained Pinus palustris (longleaf pine) stands of the southeastern United States has become an objective of land managers. Although growing‐season fire is required for successful wiregrass seed production, studies examining naturally occurring wiregrass seedling dynamics are few. This study investigates how seedling survivorship is affected by season of burn, seedling size, time since germination, and proximity to adult plants. Restoration at this research site was begun in 1992 with the planting of containerized longleaf pine and wiregrass seedlings. Study plots were established in November 1997 after a growing‐season prescribed fire (June 1996) that resulted in successful seed production and seedling recruitment. Burn treatment plots included (1) no burn (control), (2) fire in the dormant season of the first year after germination (March 1998), (3) fire in the growing season of the first year after germination (August 1998), and (4) fire in the growing season of the second year after germination (July 1999). Seedling mortality increased with growing season burning and close proximity to planted adults. Natural seedling recruitment continued into the second year after initial seed‐drop in all plots, which verifies that wiregrass seed banking occurs for a minimum of 2 years after seed drop. Where wiregrass management objectives include population expansion, seedling recruits should be allowed 1 to 2 years post‐germination without growing season fire for successful establishment.     

Ramet size equalisation in a clonal plant,Phragmites australis

The influence of shading from older generations of dead culms (standing litter) on density, growth rate and development of size structure at the ramet level was investigated in a pure stand of Phragmites australis by experimental neutral shading of plots after removal of standing litter. Initial differences in height distribution between autumn and spring cohorts disappeared in the course of shoot growth. The Gini coefficients of shoot heights and estimated shoot weights indicated that the size structure of the shoots became more equal with increasing mean size in both shaded and unshaded plots. Relative growth rate for height (RHGR) and weight of individual shoots was negatively related to shoot size during the early and presumably storage-dependent growth period, suggesting a strong support for growth of smaller shoots. No etiolation was indicated by mean or maximum height in shaded and unshaded plots, or by the relationship between shoot height and weight. Mean shoot density was significantly lower in shaded than in unshaded plots in one of two shade treatment years. A regression model indicated a small but significant effect of shoot density on the approximately linear relationship between RHGR and the logarithm of height. The growth rate of small shoots was slightly larger at low than at high shoot density. Therefore, it is suggested that the shade from standing litter in P. australis stands can decrease shoot natality in the spring cohort, and thereby increase the support to fewer small shoots.     

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.     

Testing a new hypothesis: plant vigor and phylloxera distribution on wild grape in Arizona

Summary Longer, meaning more vigorous, shoots of a wild grape clone (Vitis arizonica) were more susceptible to attack by second and third generations of leaf-galling grape phylloxera,Daktulopsphaira vitifoliae, as the growing season progressed. Although there was no significant difference in mean shoot length between attacked and unattacked shoots within a clone at the beginning of shoot elongation, attacked shoots were significantly longer than unattacked shoots when elongation had ceased (P<0.01). Also, long attacked shoots had a significantly greater population of phylloxera galls than short attacked shoots (P<0.01) as the season progressed. The phylloxera population on long shoots increased rapidly while the population on short shoots remained the same. Longer shoots also produced significantly more axillary shoots than shorter shoots as the season progressed (P<0.001), and the number of axillary shoots accounted for 66 percent of the variance in number of attacked leaves on a shoot. Experimental evidence showed that there was a significantly greater percentage of available leaves attacked on long shoots than on short shoots (P<0.05) and the leaves on long shoots generally had a greater number of galls per leaf. The relationship between shoot length and probability of attack was also tested by comparing shoots lengths of 10 attacked clones and 10 unattacked clones at a second location. Mean shoot lengths of attacked clones were significantly longer than mean shoot lengths of unattacked clones (P<0.05), and mean shoot lengths of attacked shoots within a clone were significantly longer than unattacked shoots (P<0.001). Longer shoot length accounted for 81 percent of the variance in probability of attack. The reason for this pattern of attack was that long shoots produced newly expanding leaves over a longer time during the growing season and multivoltine phylloxera require undifferentiated tissue to initiate gall formation. Patterns of attack within a shoot were characterized by an uneven distribution of galls among leaves. This was due to development time between generations and the current availability of undifferentiated tissue at times of colonization. This study supports the hypothesis that some herbivore species are favored more by vigorous plants than by stressed plants.     

Fine root production and carbohydrate concentrations of mature longleaf pine (Pinus palustris P. Mill.) as affected by season of prescribed fire and drought

The historical range of longleaf pine (Pinus palustris P. Mill) has been greatly reduced, in part, by lack of fire. Recently, the application of fire has become an accepted practice for the restoration of longleaf pine to former parts of its natural range. This study was designed to evaluate the effects of season of prescribed fire on the root growth and nonstructural carbohydrate concentrations of longleaf pine, and identify the time of year when fire has the least negative effect on longleaf pine root processes. We found that root growth was generally less on July-burned plots than on either March- or May-burned plots and we attribute these responses to the effect of fire on interaction between root processes and the soil environment. Specifically, soil water and temperature conditions may have been less favorable for root growth on the July-burned plots compared to the March- and May-burned plots. With two years of information on the seasonal dynamics of foliage production, root growth, and root carbohydrates, we determined that at our study site, optimal prescribed fire would impact tree growth less in November through March compared to other times of the year. We also observed that severe drought during the 1998 growing season was associated with a 3-month delay in peak root growth, and prolonged drought in 1999 through 2000 coincided with a reduction in root starch storage. We conclude that season of prescribed fire potentially affects root processes, but that severe or prolonged drought may either interact with or override these effects.     

Ectomycorrhizal root growth in scots pine stands in response to manipulation of litter and humus layers

 The effect on ectomycorrhizal root growth in a nitrogen-enriched planted stand of Scots pine (Pinus sylvestris L.) on podzolic sandy soil to manipulation of litter and humus layers (removal, doubling and control treatments) was examined, and compared to ectomycorrhizal root growth in an untreated naturally established Scots pine stand on nutrient-poor non-podzolic sandy soil. Half a year after manipulation of litter and humus layers in the planted stand, ingrowth-cores to a depth of 60 cm were installed in both stands. Scots pine roots were sampled four times during two growing seasons. Ectomycorrhizal roots were found at all sampled soil depths to 60 cm in all plots. Root growth and ectomycorrhizal development were greater in the naturally established stand than in all plots in the planted stand. Numbers of ectomycorrhizal root tips in the litter and humus removal plots were generally higher than in the control plots in the planted stand until May 1992. Doubling litter and humus did not significantly affect root length or the numbers of ectomycorrhizal root tips. The N _dissolved , NH₄ ⁺ and NO₃ ^– concentrations and the organic matter content in the upper 5 cm of the mineral soil in the planted stand on podzolic sandy soil were generally higher and the pH significantly lower than in the naturally established stand on non-podzolic sandy soil. Root growth and ectomycorrhizal development in the secondary stand may have been negatively affected by the chemical composition of the podzolic sandy soil. Accepted: 19 March 1997     

Stratification and seasonal stability of diverse bacterial communities in a Pinus merkusii (pine) forest soil in central Java,Indonesia

In Java, Indonesia, many nutrient-poor soils are intensively reforested with Pinus merkusii (pine). Information on nutrient cycles and microorganisms involved in these cycles will benefit the management of these important forests. Here, seasonal effects on the stratification of bacterial community structure in the soil profile of a tropical pine forest are described, and differences in bacterial communities are related to chemical and physical soil parameters. Culture-independent community profiles of litter, fragmented litter and mineral soil layers were made by denaturing gradient gel electrophoresis (DGGE) of 16S rDNA-specific polymerase chain reaction (PCR) fragments. The community profiles of the different soil layers clustered separately, correlating with significant differences in organic matter content between the three layers. The bacterial communities appeared to be stable during the wet season of 1998. The drought in 1997, caused by the El Ni?o climatic effect, did not influence the bacterial communities in fragmentation and mineral soil, although moisture content and other soil parameters were markedly lower than in the wet season. However, communities in litter were influenced by drought. In the litter layer, the moisture content was significantly lower than in the fragmentation and mineral layers during the dry season. A clone library was made from a litter sample taken during the wet season. Partial sequencing of 74 clones and linking the DGGE banding positions of these clones to bands in the DGGE profile of the sample from which the clone library was derived showed considerable bacterial diversity. Alpha-proteobacteria (40.5% of the clones, of which 57% belonged to the Rhizobium-Agrobacterium group) and high-G+C content, Gram-positive bacteria (36.5%) dominated the clone library.     

Growth and reproduction of aphids and levels of free amino acids in Scots pine and Norway spruce in an open-air fumigation with ozone

To estimate the susceptibility of conifer seedlings to aphids under future tropospheric ozone levels, Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were exposed to ambient and elevated ozone levels in an open-air exposure system. Growth and reproduction of the aphids Schizolachnus pineti and Cinara pinea on Scots pine and Cinara pilicornis on Norway spruce were monitored. Levels of free amino acids in foliage and young shoots were used as indicators of host plant quality. In elevated treatment plots the ozone doses were between 1.2 and 1.7 times the dose in ambient plots in 1990–93. Half of the seedling material in 1992–93 was subjected to nitrogen fertilization treatment to evaluate the effects of increased N deposition. In 1990, population density of S. pineti on pine did not differ between ambient and elevated ozone treatments during growing season, but remained higher in the elevated ozone plot than in the ambient plot at the end of the growing season. This was associated with elevated levels of glutamic acid in foliage. In August 1992, the numbers of S. pineti were consistent between the two ambient ozone plots, but deviated highly between the two ozone-fumigated plots. Glycine concentration in pine foliage was elevated by ozone, but free amino acid concentrations were not related to aphid performance. In 1993, ozone and nitrogen did not significantly affect the relative growth rate (RGR) of S. pineti or C. pinea nymphs on Scots pine, but glutamic acid concentration in foliage was increased by nitrogen fertilization. On Norway spruce, fecundity of C. pilicornis females was higher in elevated ozone treatment, but RGR of nymphs was not affected in 1992. In 1993, RGR of C. pilicornis nymphs was increased by nitrogen fertilization in June, but not affected by ozone. Nitrogen fertilization increased the levels of total free amino acids, aspartic acid, glutamic acid and proline in elongating shoots of Norway spruce, and ozone reduced the concentrations of valine and γ-butyric acid. Our results suggest that availability of nitrogen from soil has a stronger impact on the concentrations of free amino acids in conifer seedlings than ozone. Some episodes of high ozone concentration may increase free amino acids in foliage. Aphid response to ozone was extremely variable, in agreement with previous laboratory experiments. The expected 20–70% increase in ambient concentrations of tropospheric ozone may in some occasions enhance aphid performance on Scots pine and Norway spruce seedlings, but in most cases the ozone effect on the susceptibility of conifer seedlings to sucking insect pests is not important.     

Responses of root‐crown bearing shrubs to differences in fire regimes in Pinus palustris (longleaf pine) savannas: Exploring old‐growth questions in second‐growth systems

Question: What are the effects of fire season and intensity on resprouting of different root‐crown bearing shrub species in second‐growth Pinus palustris (longleaf pine) savannas? Location: northern Florida and eastern Louisiana, USA. Methods: In Florida, quadrats were burned biennially either during the dormant season or the growing season. In Louisiana, we applied intensity treatments to quadrats by manipulating ground‐cover fuels, just prior to biennial growing season fires. Maximum fire temperatures were measured, and stem densities were censused before and after fires in both regions. Results: After dormant season fires in Florida, stem densities were seven times greater than initial levels for Hypericum spp. In contrast, growing season fires reduced densities of H. brachyphyllum by 65%, but did not change densities of H. microsepalum. Only resprouting of H. microsepalum decreased with increased fire intensity. In Louisiana, fire intensity influenced Ilex vomitoria, but not Quercus spp. Following fires, stem densities oil. vomitoria were five times greater in fuel removal than fuel addition areas. Conclusions: Past use of dormant season fires likely contributed to increased abundances of some species of root‐crown bearing shrubs observed today in old‐growth savannas. Reintroduction of growing season fires will be effective in maintaining or decreasing stem densities, depending on species and fuel type. Genet mortality and stem density reductions appear most likely in areas at localized scales where tree falls and needle coverage create hotspots in Pinus palustris savannas.     

Seasonal patterns of partitioning and remobilization of 14C in the invasive rhizomatous perennial Japanese knotweed (Fallopia japonica (Houtt.) Ronse Decraene)

Resource partitioning between shoot growth, storage and reproduction is poorly understood in many clonal plant species. This study documents seasonal patterns of growth, ¹⁴C-labelled photoassimilate distribution and remobilization in the invasive rhizomatous species Fallopia japonica (Japanese knotweed). Biomass accumulation above- and below-ground in F. japonica was rapid. By September, rhizome biomass had increased 18-fold from the initial harvest in May (representing 48% of total plant biomass) and this was maintained over winter. Patterns of ¹⁴C allocation from F. japonica shoots labelled at different times of year show that as the season progressed, the rhizomes became an increasingly important sink for current assimilate (the percentage of ¹⁴C recovered from rhizomes was 35% in August and 67% in September) and the corresponding retention of assimilate by established shoots declined. The percentage of ¹⁴C exported to roots was greatest in August. Relatively little photoassimilate was exported to other shoots on the plant, or to flowers. Recycling of photoassimilate was fairly tight in this species and ¹⁴C fixed by shoots in early May 1999 or September 1999 was remobilized to the rhizome prior to shoot senescence and death. Some of this ¹⁴C was then remobilized to new shoots early the following spring. These characteristics may contribute to the success of F. japonica in colonizing a variety of contrasting habitats, often with serious management implications.     

Effects of litter removal on arthropod communities in pine plantations

Natural and anthropogenic disturbances can cause abrupt changes in trophic interactions by altering the rate, timing, or composition of organic inputs to ecological systems which in turn can shift patterns of species dominance. We examined the short-term effects of litter removal on soil fauna in pine plantations of three different species (longleaf, Pinus palustris; loblolly, P. taeda; and slash, P. elliottii) using a manipulative experiment, with the goal of examining differences among dominant orders of arthropods and differences among timber types. We sampled arthropods once per month for 6 months immediately following raking, and found that removal of the litter caused significant changes to abundance or presence of five of the nine dominant orders. Reductions in abundances of arthropod orders were most apparent in loblolly pine stands, while increases in abundance were more common in longleaf and slash pine stands. The differential impact among orders suggests that removal of the litter layer is likely to alter trophic interactions by changing the relative abundance of functional groups. Repeated litter removal via raking could have negative repercussions on ecosystem stability. Finally, nutrient additions through fertilization seem unlikely to mitigate the changes imposed on the arthropod community through litter removal.     

Pine savanna overstorey influences on ground‐cover biodiversity

Question: Does the overstorey of pine savannas influence plant species biodiversity in the ground cover? Location: Camp Whispering Pines (30°41’N; 90°29’W), eastern Louisiana (USA). Methods: We used ecologically sensitive restoration logging to remove patches of Pinus palustris (longleaf pine) in a second‐growth loess plain Pinus palustris savanna managed using frequent lightning season fires. Five years later, we measured numbers of vascular plant species and transmitted light in replicated 100‐m² plots. Treatments involved three different overstorey conditions: no overstorey for 5 years, no overstorey for several decades, and overstorey pines present for decades. Results: Both recent and long‐term openings contained, on average, about 100 vascular plant species per 100 m², 20% more than in similar‐sized areas beneath overstorey trees. Responses varied with life form; more herbaceous species occurred in recent and older overstorey openings than beneath overstorey trees. Total numbers of all species and of less abundant forb species were positively and linearly related to light transmitted to ground level. Those species responding to openings in the overstorey and positively associated with increased transmitted light levels were monocarpic and shortlived perennial forb and grass species with a seed bank in the soil. In addition, community structure, as reflected in species composition and abundances, appeared to vary with canopy condition. Conclusions: Restoration involving ecologically sensitive removal of patches of overstorey pines in frequently burned pine savannas should benefit the ground cover and increase plant species biodiversity as a result of increased abundance of seed bank species.     

Experimental demography of rhizome populations of establishing clones of Solidago altissima

1 Solidago altissima forms large compact below-ground rhizome systems. The rhizome systems of 30 genets in a population that had colonized an old-field in 1984 were mapped yearly in situ from 1988 to 1992. Branching angles, lengths and survival of rhizomes, and production of daughter rhizomes, were calculated from digitized maps.
2 We hypothesized that genets of S. altissima would prevent intraclonal crowding by producing longer but fewer rhizomes as they increased in size. In addition, we tested the effects of environmental factors on rhizome dynamics by (i) mowing in late summer, (ii) cutting of rhizome connections, and (iii) removal of close neighbour plants.
3 Most rhizomes were initiated at an angle of 60°−65° relative to the parent rhizome. Subsequent growth tended towards the parent axis and towards more centrifugal growth. Rhizome angles were not influenced by any treatment.
4 In control genets, rhizome density increased 1.3-fold from March 1988 to March 1992. The number of newly produced rhizomes per genet decreased and their length increased from 1988 to 1992. Longer rhizomes had a higher survival probability than shorter rhizomes did. The number of newly produced rhizomes per parent rhizome was positively correlated with mean rhizome length per genet.
5 Mowing strongly decreased rhizome density from 1988 to 1992, mainly due to reduced production of new rhizomes.
6 Rhizome cutting led to a slight decline in the rhizome population and to shorter lengths of rhizomes in their second year compared with controls.
7 The removal of close neighbour plants led to increased rhizome densities. These rhizomes grew longer than controls in unmown plots and shorter in mown plots and had higher probabilities of surviving into their second year.
8 Yearly mowing could successfully prevent the further spread of S. altissima because it acts on rhizome and shoot dynamics as well as on seed output.     

Interactions Between Shoot Age Structure, Nutrient Availability and Physiological Integration in the Giant Bamboo Phyllostachys pubescens

Abstract: The age structure of adult shoots, the nutrient availability of the habitat, and their interaction, are important factors influencing the productivity of bamboo groves. In a field fertilization experiment over two years we examined the impact of physiological integration on the emergence, growth, and survival of new shoots of Phyllostachys pubescens, a giant woody bamboo. Impacts of physiological integration were compared using uniform and patchy fertilization treatments. The number of new shoots emerging per plot significantly increased with the application of fertilizer (NPK), but the increase was constrained by the age structure of adult shoots. In a year with a high proportion of shoots with new (first-year) leaves many more new shoots emerged than in a year with a low proportion of new leaves. Mean survivorship of the new shoots was constant at 20 % for all treatments in both years of study. Surprisingly, fertilization did not increase DBH or height of the new shoots, suggesting that shoot size was fixed early in ontogeny. The pattern of fertilizer application, either uniform or patchy, did not affect the total number of new shoots produced. In the case of patchy application, more new shoots developed in unfertilized patches in comparison to uniformly unfertilized plots, probably because these parts of the clone received resources via the rhizomes from the adult shoots in adjacent fertilized patches. The production of new shoots in fertilized patches, in turn, was lower than that in uniformly fertilized plots as a result of this physiological integration. The results are discussed in the general context of the impact of integration on clonal plant performance in dense stands.     

鼎湖山退化马尾松林、混交林和季风常绿阔叶林土壤全磷和有效磷的比较

土壤全磷和有效磷浓度的变化随林型和季节不同而异,总的来说,其大小顺序为:季风常绿阔叶林> 混交林>马尾松林(林型);夏季>冬季>春季>秋季(季节),有效磷浓度为:秋季>春季>冬季>夏季(季 节)。收割林下层和凋落物这种人为干扰活动对土壤全磷含量的影响不明显,但对土壤有效磷含量具有显著 的影响。在试验开始时(1990年5月),土壤全磷和有效磷浓度在马尾松林保护样地(停止人为干扰)和处理样 地(按当地习惯继续收割林下层和凋落物)间的差异均不显著。经七年多的试验后,土壤全磷浓度在保护样地 和处理样地间仍十分相似,但土壤有效磷浓度在保护样地显著高于处理样地,说明保护样地在停止人为干扰 后相对于继续受干扰的处理样地其林地条件得到了改善,从而使土壤有效磷含量也得到提高。只要停止人为 干扰,鼎湖山退化马尾松林土壤有效磷供应力是可以自然恢复到季风常绿阔叶林的水平。     

The role of litter in an old-field community: impact of litter quantity in different seasons on plant species richness and abundance

Summary We studied the effect of removing and adding plant litter in different seasons on biomass, density, and species richness in a Solidago dominated old-field community in New Jersey, USA. We removed all the naturally accumulated plant litter in November (658 g/m²) and in May (856 g/m²) and doubled the amount of litter in November and May in replicated plots (1 m²). An equal number of plots were left as controls. Litter removal and addition had little impact on total plant biomass or individual species biomass in the growing season following the manipulations. Litter removal, however, significantly increased plant densities but this varied depending upon the season of litter removal, species, and life history type. Specifically, the fall litter removal had a much greater impact than the spring litter removal suggesting that litter has its greatest impact after plant senescence in the fall and prior to major periods of early plant growth in spring. Annual species showed the greatest response, especially early in the growing season. Both spring and fall litter removal significantly increased species richness throughout the study. Litter additions in both spring and fall reduced both plant densities and species richness in June, but these differences disappeared near the end of the growing season in September. We concluded than in productive communities where litter accumulation may be substantial, litter may promote low species richness and plant density. This explanation does not invoke resource competition for the decline in species richness. Finally, we hypothesize that there may be broad thresholds of litter accumulation in different community types that may act to either increase or decrease plant yield and diversity.