Soil translocation by pocket gophers in a Mima moundfield

Summary We measured soil translocation due to the tunneling of valley pocket gophers (Thomomys bottae) in a Mima moundfield at Miramar Mounds National Landmark, San Diego, California, from December, 1984 through December, 1985. We placed 1-l soil plugs containing 20 11-g iron pellets into pocket gopher tunnels at locations between mound tops and points about one mound radius beyond mound edges. After about 4–10 d, sites to which the marker-containing soil had been translocated were located with a metal detector and the horizontal and vertical displacements measured. Between 1 October and 15 May (the cooler, wetter portion of the year), pocket gophers removed an average of 63% of the experimental plugs and moved an average of 38% of the markers that we recovered. From 15 May through 1 October (the hotter, drier portion of the year), only 32% of plugs were cleared and 12% of the recovered markers were moved. On average, markers that were moved were displaced 41 cm moundward and 4.9 cm upward in elevation. The intensity of moundward translocation increased with distance from the mound center. At a distance of 0.5–1.0 mound radius beyond the edge of the mound, the moundward translocation tendency averaged 71 cm. The intensity of moundward translocation was also inversely related to maximum mound height. These observations provide strong support for the fossorial rodent hypothesis of Mima mound origin, and constitute a first step in development of a mathematical model of mound formation.     

The urban fire ant paradox: native fire ants persist in an urban refuge while invasive fire ants dominate natural habitats

In contrast to the widespread extirpation of native fire ants (Solenopsis geminata) across southern US following the invasion by imported red fire ants (S. invicta), some residential areas of Austin form unexpected refuges for native fire ants. Ironically, these urban environments provide refuges for the native fire ants while adjacent natural habitats have been overrun by invasive fire ants. Resistance to invasive fire ants in these urban areas occurs mainly in older residential properties constructed prior to the S. invicta invasion, while more recent construction has allowed establishment by S. invicta. The invasive ability of S. invicta is often attributed to escape from parasitoids and efficient dispersal of polygyne multiple queen colonies. Here we also show the importance of landscape parameters in the invasion process, where low levels of disturbance and continuous plant cover in older residential areas form possible barriers to colonization. Dense leaf cover (high NDVI) was also found to be associated with native ant refuges. Long term residential land ownership may have resulted in lower recent disturbance levels and increased plant cover that support refuges of native fire ants.     

Cophylogeny and disparate rates of evolution in sympatric lineages of chewing lice on pocket gophers

Although molecular-based phylogenetic studies of hosts and parasites are increasingly common in the literature, no study to date has examined two congeneric lineages of parasites that live in sympatry on the same lineage of hosts. This study examines phylogenetic relationships among chewing lice (Phthiraptera: Trichodectidae) of the Geomydoecus coronadoi and Geomydoecus mexicanus species complexes and compares these to phylogenetic patterns in their hosts (pocket gophers of the rodent family Geomyidae). Sympatry of congeneric lice provides a natural experiment to test the hypothesis that closely related lineages of parasites will respond similarly to the same host. Sequence data from the mitochondrial COI and the nuclear EF-1alpha genes confirm that the two louse complexes are reciprocally monophyletic and that individual clades within each species complex parasitize a different species of pocket gopher. Phylogenetic comparisons reveal that both louse complexes show a significant pattern of cophylogeny with their hosts. Comparisons of rates of nucleotide substitution at 4-fold degenerate sites in the COI gene indicate that both groups of lice have significantly higher basal mutation rates than their hosts. The two groups of lice have similar basal rates of mutation, but lice of the G. coronadoi complex show significantly elevated rates of nucleotide substitution at all sites. These rate differences are hypothesized to result from population-level phenomena, such as effective population size, founder effects, and drift, that influence rates of nucleotide substitution.     

Patterns of size change in late Neogene pocket gophers from the Meade Basin of Kansas and Oklahoma

A dense late Neogene history of pocket gophers from the Meade Basin of south-western Kansas and north-western Oklahoma preserves a detailed record of size change, as indicated by mean length of the lower fourth premolar. A ‘time for space substitution’ interpretation results in the tentative recognition of a pattern analogous to character displacement, in which the small to medium-sized Pliogeomys buisi, P. louderbachi, Geomys minor and G. floralindae are considered ecological analogues and treated as a single ecological entity, Avatar A. The larger G. jacobi and G. quinni are combined as Avatar B. Statistical analyses confirm that populations of Avatar A are of equal size when allochronic from Avatar B, and significantly smaller when synchronic (and sympatric) with Avatar B. However, this pattern is complicated by the observation that Avatar A began dwarfing before the appearance of Avatar B. This historical perspective suggests that static size patterns among modern taxa may have a complex history. Occasional diminutive geomyid transients (G. adamsi, Thomomys cf talpoides) are always paired with significantly larger, medium-sized Geomys. Although a random walk explanation for the pattern of size changes is unlikely, we are currently unable to distinguish between environmental change and competition as the likely forcing mechanisms.     

Morphology, genetics, and ecology of pocket gophers (genus Geomys) in a narrow hybrid zone

A population of hybrid pocket gophers ( Geomys bursarius × G. lutescens ) exists 1–2 km west of Oakdale, Antelope County, Nebraska, U.S.A. The hybrids occur in soil that has characteristics intermediate between that occupied by G. lutescens (sand) and by G. bursarius (silt loam); the vegetation associations on the different soils are Sandhills Prairie and Tall-grass Prairie, respectively, with mixed prairie on the intermediate soils. Hybrids are identifiable on the basis of both qualitative and quantitative morphological characteristics, allozymes and karyology. Concordance between morphological, allozymic, karyotypic and ecological data sets is very high. Hybrids appear to reproduce normally and survive well; i.e. they suffer no obvious loss of fitness. Backcrossing to either parental type is apparently rare. The parental species each support obligate parasitic lice ( Geomydoecus : Trichodectidae) of different species; these species are not sister species. We suggest that hybrid zones resulting from primary and secondary contact may be distinguished by (1) concordance of clines in different character sets, (2) fossil and biogeographic data, and (3) parasite data. We conclude that this zone resulted from secondary contact, and that the zone is maintained either by selection against hybrids (less likely) or by hybrid superiority (more likely).     

The effects of fire on scarlet gilia: an alternative selection pressure to herbivory?

Summary The primary goals of this paper were to experimentally assess the relative importance of fire, a potentially important selective agent involved in shaping scarlet gilia's compensatory response and, in general, determine scarlet gilia's response to fire. Burn treatment results demonstrated that fire was not an important selective agent involved in shaping scarlet gilia's compensatory response. The most common response to fire was the production of one or more new clonally derived rosettes. This was an unexpected result; this typically monocarpic herb rarely produces clonal offspring. Although rosette production lessened the detrimental effects of fire by giving plants that cloned a second chance to flower, these newly formed rosettes delayed flowering for at least one year and had significantly higher overwinter mortality rates than rosettes from unburned control plots. In addition, significantly fewer individuals from the burn treatments flowered and there was significantly higher immediate mortality. There was, however, no detrimental effect on the reproductive success (seed production) of individuals that flowered following the burn. Overall, cumulative estimates of plant performance suggest that at the population level fire results in a 4.5-fold decrease in relative plant fitness. However, fire-induced seed germination could negate the detrimental effects of fire on the population dynamics of scarlet gilia. In the year of the burn there was a 116-fold increase in the number of germinating seeds and by the second year this translated into an approximate 6-fold difference in the number of surviving rosettes. Two alternative candidates, frost damage and ungulate trampling, can cause the removal of apical dominance and a response similar to that generated by ungulate and insect herbivores. However, they are probably minor factors favoring selection toward growth compensation; experimental and observational results deomonstrate that apical dominance was removed in only 3% of plants exposed to freezing temperatures and ungulate trampling caused breakage and release of apical dominance in only 0.2% of plants.     

The relationship between fire history and an exotic fungal disease in a deciduous forest

Exotic diseases have fundamentally altered the structure and function of forest ecosystems. Controlling exotic diseases across large expanses of forest has proven difficult, but fire may reduce the levels of diseases that are sensitive to environmental conditions. We examined Cornus florida populations in burned and unburned Quercus–Carya stands to determine if burning prior to anthracnose infection has reduced the impacts of an exotic fungal disease, dogwood anthracnose, caused by Discula destructiva. We hypothesized that fire has altered stand structure and created open conditions less conducive to dogwood anthracnose. We compared C. florida density, C. florida health, and species composition and density among four sampling categories: unburned stands, and stands that had burned once, twice, and 3 times over a 20-year period (late 1960s to late 1980s). Double burn stands contained the greatest density of C. florida stems (770 stems ha⁻¹) followed by triple burn stands (233 stems ha⁻¹), single burn stands (225 stems ha⁻¹) and unburned stands (70 stems ha⁻¹; P < 0.01). We observed less crown dieback in small C. florida trees (<5 cm diameter at breast height) in burned stands than in unburned stands (P < 0.05). Indicator species analysis showed that burning favored species historically associated with Quercus–Carya forests and excluded species associated with secondary succession following nearly a century of fire suppression. Our results suggest that fire may mitigate the decline of C. florida populations under attack by an exotic pathogen by altering forest structure and composition. Further, our results suggest that the burns we sampled have had an overall restorative effect on forest communities and were within the fire return interval of the historic fire regime. Consequently, prescribed fire may offer a management tool to reduce the impacts of fungal disease in forest ecosystems that developed under historic burning regimes.     

Patch to landscape patterns in post fire recruitment of a serotinous conifer

Obligate seeding species are highly specialized to fire disturbance and many conifers such as cypress, which are adapted to high intensity stand-replacing fires, have canopy seed banks stored in serotinous cones. Resilience of these trees to fire disturbance is a function of disturbance frequency and one focus of this study was to determine the effect of patch age on postfire recruitment. A second focus was to determine the extent to which fire induced a landscape level change in the location of the forest boundary. Prior to a fire in 1994, a large Cupressus sargentii forest was a mosaic landscape of different aged patches of nearly pure cypress bordered by chaparral. Patches less than 60 years of age were relatively dense with roughly one tree every 1–2 m² but older patches had thinned to one tree every 3–15 m². Older trees had substantially greater canopy cone crops but the stand level seed bank size was not significantly correlated with stand age. Fire-dependent obligate seeding species are sensitive to fire return interval because of potential changes in the size of seed banks – facing both a potential `immaturity risk' and a `senescence risk'. At our site, C. sargentii regeneration was substantial in stands as young as 20 years, suggesting that fire return interval would need to be shorter than this to pose any significant risk. Reduced seedling recruitment in stands nearly 100 years of age may indicate risk from senescence is greater, however, even the lowest density seedling recruitment was many times greater than the density of mature forests – thus this cypress would appear to be resilient to a wide range of fire return intervals. Changes in landscape patterning of forest and chaparral are unlikely except after fire. Factors that inhibit tree establishment within the shrubland, as well as factors that affect shrub establishment within the forest border likely affect the `permeability' of this ecotone. After the 1994 fire this boundary appeared to be stable in that cypress recruited best within the shadow of burned canopies and cypress were weak invaders of adjacent shrublands.     

Effects of controlled livestock grazing and annual prescribed fire on epigeal termite mounds in a savannah woodland in Burkina Faso

The diversity and abundance of epigeal termite mounds were investigated in response to controlled livestock grazing and annual prescribed fire in a Sudanian savannah-woodland in Tiogo State Forest. Sampling of termite mounds was carried out in 4×4 subplots of 0.25 ha in a split-plot experimental design during the rainy season in 2002. There were two main plots of which one was fenced to exclude livestock grazing and the second exposed to grazing. Each of the main plots included 4 subplots with annual prescribed fire since 1992 and 4 subplots without fire. Data were collected on the number and characteristics of termite mounds. A mean density of 698 mounds ha⁻¹ was recorded. Mounds built by Trinervitermes spp. were the most abundant followed by Cubitermes spp., Macrotermes subhyalinus and M. bellicosus. The large mound-builders Macrotermes spp. dominated the community interms of basal area (96% of the total) and above-ground volume (99%). The diversity of mound types was notaffected by livestock grazing and annual early prescribed fire (P>0.05). There was no statistical effect of livestock grazing on mound density, whereas a strong depressive effect of annual fire was observed for Trinervitermes spp. mound density (P=0.012). In this ecosystem, annual prescribed fire appeared to be the major determinant for termite mound abundance. Received 2 February 2007; revised 23 October 2007 and 21 January 2008; accepted 22 Feburary 2008.     

The effects of neighbors on the growth and survival of shrub seedlings following fire

Where plant species vie for limited resources, disturbances might preclude competition by releasing a flush of nutrients, or by reducing biomass and thereby diminishing the consumption of resources. However, if new seedlings colonize in clumps, they may still deplete resources within the local aggregations, which may then reduce their growth and survivorship. We investigated competition among seedlings in a burned area by examining the relationship between the performance of newly established shrub seedlings of Ceanothus impressus and (1) the proximity and (2) the identity of their near neighbors. We also investigated the relationship between neighbor proximity and the availability of water. Both survivorship and growth of C. impressus were positively associated with increasing distance to near neighbors, in a manner consistent with resource competition. The availability of water (as determined by pre-dawn xylem pressure potentials) tended to be greater when neighbors were farther away, providing evidence that water was a resource for which plants were competing. This conclusion is reinforced by the finding that the effects of neighbors were stronger in drought years, suggesting that yearly variation in the availability of an important resource (water) can affect the strength of competitive interactions. This suggests that after disturbances, when some resources are apparently abundant on a large scale, competition may be important in determining the small scale patterns of seedling growth and survival.     

Potential cause of lethality of an allele implicated in social evolution in fire ants

The gene Gp-9 is believed to have a major effect on colony social organization in fire ants, with the presence of b-like alleles in a colony associated with multiple-queen (polygyne) organization. Queens and workers of polygyne Solenopsis invicta homozygous for the b-like allele designated b suffer reduced viability compared to other genotypes, and bb queens do not survive to become egg-layers. Thus, the b allele effectively acts as a recessive lethal. This allele differs from the remaining b-like alleles (designated b′), as well as all other Gp-9 alleles, by encoding a lysine at position 151 in the protein product, suggesting that this substitution is responsible for its deleterious effects. We tested this hypothesis by comparing frequencies of b′b′ and bb homozygotes, first in queens of Solenopsis richteri and S. invicta, then in S. invicta workers from populations polymorphic for the two b-like alleles. We found that almost 20% of S. richteri queens were b′b′ homozygotes, compared to the virtual absence of bb homozygotes among S. invicta queens, and that 5–18% of S. invicta workers bore genotype b′b′, compared to the apparent lack of bb workers in the same populations. While we cannot entirely rule out involvement of other genes in complete gametic disequilibrium with Gp-9, our data are consistent with the hypothesis that the Lys¹⁵¹ residue in GP-9 protein confers the deleterious effects of the b allele in homozygous condition, possibly by impairing the protein’s function through interference with ligand binding/release or hindrance of dimer formation.     

Temporal and spatial patterns in fire occurrence during the establishment of mixed‐oak forests in eastern North America

Question: What was the role of fire during the establishment of the current overstory (ca. 1870–1940) in mixed‐oak forests of eastern North America? Location: Nine sites representing a 240‐km latitudinal gradient on the Allegheny and Cumberland Plateaus of eastern North America. Methods: Basal cross‐sections were collected from 225 trees. Samples were surfaced, and fire scars were dated. Fire history diagrams were constructed and fire return intervals were calculated for each site. Geographic patterns of fire occurrence, and fire‐climate relationships were assessed. Results: Fire was a frequent and widespread occurrence during the formation of mixed‐oak forests, which initiated after large‐scale land clearing in the region ca. 1870. Fire return ranged from 1.7 to 11.1 years during a period of frequent burning from 1875 to 1936. Fires were widespread during this period, sometimes occurring across the study region in the same year. Fires occurred in a variety of climate conditions, including both drought and non‐drought years. Fires were rare from 1936 to the present. Conclusions: A variety of fire regime characteristics were discerned. First, a period of frequent fire lasted approximately 60 years during the establishment of the current oak overstory. Second, fire occurred during a variety of climate conditions, including wet climates and extreme drought. Finally, there was within‐site temporal variability in fire occurrence. These reference conditions could be mimicked in ongoing oak restoration activities, improving the likelihood of restoration success.     

Three centuries of fire in montane pine‐oak stands on a temperate forest landscape

Question: What was the role of fire in montane pine‐oak (Pinus‐Quercus) stands under changing human land uses on a temperate forest landscape in eastern North America? Location: Mill Mountain in the central Appalachian Mountains, Virginia, US. Methods: A dendroecological reconstruction of fire history was generated for four stands dominated by xerophytic pine and oak species. The fire chronology began under presettlement conditions following aboriginal depopulation. Subsequent land uses included European settlement, iron mining, logging, and US Forest Service acquisition and fire protection. Results: Fires occurred approximately every 5 years until 1930 without any evidence of a temporal trend in fire frequency. Burning ceased after 1930. Area‐wide fires affecting multiple pine stands were common, occurring at intervals of approximately 16 years. Most living pines became established during the late 1800s and early 1900s. Dead pines indicated that an older cohort established ca. 1730. Most hardwoods were established between the 1920s and 1940s. Conclusions: Except for fire protection, changes in land use had no discernible influence on fire frequency. Lightning ignitions and/or large fire extent may have been important for maintaining frequent burning in the 1700s, while fuel recovery may have constrained fire frequency during later periods. The disturbance regime appears to be characterized by frequent surface fires and occasional severe fires, insect outbreaks or other disturbances followed by pine recruitment episodes. Industrial disturbances appear to have had little influence on the pine stands. The greatest impact of industrial society is fire exclusion, which permitted hardwood establishment.     

华北低丘山区栓皮栎生态系统氧同位素日变化及蒸散定量区分

利用稳定同位素技术对华北低丘山区栓皮栎生态系统氧同位素日变化及蒸散定量区分进行研究,为华北低丘山区森林生态系统水汽交换研究提供基础。试验采用离轴积分腔输出光谱技术(OA-ICOS)连续测定生态系统不同高度水汽浓度和δ18O值,同时采用真空提取和液态水同位素分析仪测定枝条和土壤的δ18O值。结果显示,4个晴天中大气水汽浓度日变化复杂,变化趋势差异大,而δ18O日变化均成高-低-高的"V"型变化,最小值出现在12:00—18:00。Keeling方程在10:00—12:00的相关系数R2均大于0.71,方程达到极显著水平,表明此时段蒸腾速率较高,满足植物蒸腾的同位素稳定态假设。利用Keeling方程估算的栓皮栎生态系统δET值有相似的低-高-低日变化,与大气的δv值变化趋势相反。同位素分割结果显示栓皮栎生态系统中蒸腾占蒸散比例日变化呈现低-高-低的趋势,10:00—14:00蒸腾占蒸散比例达到90%以上,尽管6:00—10:00和14:00—18:00的蒸腾占蒸散比例下降,但平均值仍高达69.38%,表明华北低丘山区栓皮栎生态系统的蒸散主要来源于植物蒸腾。     

Interactions between fire,mycophagous mammals,and dispersal of ectromycorrhizal fungi in Eucalyptus forests

Several species of marsupials in Eucalyptus forests in Australia feed predominantly on the sporocarps of hypogeous fungi. This feeding is apparently beneficial to the fungi as it results in dispersal of spores. As these fungi are in almost all cases ectomycorrhiza-forming species, mycophagy by mammals may play an important role in the maintenance of mycorrhizal symbiosis in Eucalyptus forests. Fire is frequent and a dominant ecological factor in these forests, and this study tested the hypothesis that fire triggers both increased sporocarp production by some hypogeous ectomycorrhizal fungi associated with eucalypts, and increased mycophagy by mammals. Three experimental burns were set in E. tenuiramus forest in southeastern Tasmania. Digging activity (which reflects feeding on hypogeous fungi) by a mycophagous marsupial, the Tasmanian bettong Bettongia gaimardi, increased up to ten-fold after fire, with a peak about 1 month post-fire. This was associated with a similar pattern of increase in sporocarp production, which was due to species in the family Mesophelliaceae (especially Castoreum tasmanicum and Mesophellia spp.). This family appears to have radiated in association with eucalypts and has an exclusively Australasian distribution, unlike many of the other ectomycorrhizal fungi collected in this study which are cosmopolitan and have broad host ranges. No B. gaimardi were killed by fire, and there was no increase in mortality following fire. Population density increased after fire as a result of immigration of adult males. However, body condition and fecundity of individual B. gaimardi were maintained at pre-fire levels. This suggests that the availability of energy to B. gaimardi increased as a result of fire, and the fact that the contribution of fungus to the diet of B. gaimardi was high on burnt relative to control sites suggests further that this increase in energy availability was provided by hypogeous fungi. Effects of fire on hypogeous fungi and B. gaimardi were short-lived; all measured variables returned to control values about 4 months after fire. The capacity of B. gaimardi to survive fire and to harvest the increased sporocarp production triggered by fire provides a mechanism for the rapid dispersal of spores after fire. This should result in the establishment of ectomycorrhizae very early in post-fire succession. Because only some species of ectomycorrhizal fungi fruited in response to burning, fire probably has a strong influence on community structure among ectomycorrhizal fungi.     

Population and clonal level responses of a perennial grass following fire in the northern Chihuahuan Desert

Relationships involving fire and perennial grasses are controversial in Chihuahuan Desert grasslands of southern New Mexico, USA. Research suggests that fire delays the resprouting of perennial grasses well after two growing seasons. However, such results are confounded by livestock grazing, soil erosion, and drought. Additionally, post-fire grass responses may depend on initial clone size. We evaluated the effects of fire, grazing, and clone size on Bouteloua eriopoda (black grama) in southern New Mexico grasslands. Four 2-ha plots were established in each of four sites. Fire and grazing were applied or not applied in 1999 such that four treatment combinations were assigned randomly to plots within each site. Within each plot, small (0–10 cm² basal area), medium (10–30 cm²), and large ( > 30 cm²) clones were initially mapped in five 0.91-m² quadrats where grass attributes and litter cover were evaluated before and at the end of two growing seasons following fire. Maximum fire temperature was also measured. At a population level, canopy and litter cover were each approximately 50% less in burned than unburned areas. However, compared to initial levels, canopy height had increased by 10% at the end of the study, regardless of fire. At a clonal level, basal cover reductions were attributed mostly to large clones that survived fire. Smaller clone densities had decreased by as much as 19% in burned compared to unburned areas, and fire reduced the basal cover of medium clones. Basal and canopy cover, recruitment, and clone basal area decreased with increased fire temperatures. Almost all responses were independent of grazing, and interactive effects of grazing and fire were not detected. Fire did not kill all perennial grass clones, regardless of size. However, rapid responses were likely influenced by above-average precipitation after fire. Future studies in desert grasslands should examine how perennial grass dynamics are affected by fire, precipitation patterns, and interactions with grazing.     

Plant functional group responses to fire frequency and tree canopy cover gradients in oak savannas and woodlands

Questions: How do fire frequency, tree canopy cover, and their interactions influence cover of grasses, forbs and understorey woody plants in oak savannas and woodlands? Location: Minnesota, USA. Methods: We measured plant functional group cover and tree canopy cover on permanent plots within a long‐term prescribed fire frequency experiment and used hierarchical linear modeling to assess plant functional group responses to fire frequency and tree canopy cover. Results: Understorey woody plant cover was highest in unburned woodlands and was negatively correlated with fire frequency. C4‐grass cover was positively correlated with fire frequency and negatively correlated with tree canopy cover. C3‐grass cover was highest at 40% tree canopy cover on unburned sites and at 60% tree canopy cover on frequently burned sites. Total forb cover was maximized at fire frequencies of 4–7 fires per decade, but was not significantly influenced by tree canopy cover. Cover of N‐fixing forbs was highest in shaded areas, particularly on frequently burned sites, while combined cover of all other forbs was negatively correlated with tree canopy cover. Conclusions: The relative influences of fire frequency and tree canopy cover on understorey plant functional group cover vary among plant functional groups, but both play a significant role in structuring savanna and woodland understorey vegetation. When restoring degraded savannas, direct manipulation of overstorey tree canopy cover should be considered to rapidly reduce shading from fire‐resistant overstorey trees. Prescribed fires can then be used to suppress understorey woody plants and promote establishment of light‐demanding grasses and forbs.     

Ecophysiology of Zigadenus nuttallii,a toxic spring ephemeral in a warm season grassland: effect of defoliation and fire

Summary Zigadenus nuttallii, a highly toxic spring ephemeral in tallgrass prairie, was studied in 1985 to ascertain: 1) several ecophysiological characteristics of the species, 2) seasonal patterns of biomass accumulation, and 3) its response to defoliation and fire. The maximum photosynthetic rate of Z. nuttallii measured in unburned prairie was 13.2 moles CO₂ m^-2 s^-1 which occurred at 24–28° C and an incident quantum flux of 0.8–1.0 mmoles m^-2 s^-1. Maximum stomatal conductance measured was 5.4 mm s^-1. Early in the season, belowground storage organs (bulbs) decreased in mass and supplied much of the energy for growth of leaves, even though CO₂ uptake was possible. Buld mass did not increase until about 6 weeks after shoot emergence implying that, at this time, leaves had become a source rather than a sink for carbohydrates. The result of a single, severe defoliation event was a decrease in biomass of bulbs, leaves and reproductive structures in Z. nuttallii. Intrinsic compensatory mechanisms were not detected. In contrast, fire, which also defoliated plants, did not result in any biomass decrease at the end of the season. Improved post-fire microclimate and increased nutrient supply (extrinsic factors) may have contributed to higher photosynthetic rates and led to biomass compensation in burned prairie. These data support arguments that intrinsic compensatory mechanisms have evolved in response to chronic herbivory.     

The effects of phosphorus nutrition and soil pH on the growth of some ancient woodland indicator plants and their interaction with competitor species

The forest under-storey herbs Anemone nemorosa, Lamiastrum galeobdolon and Veronica montana are generally considered indicator species of old, broadleaved woodland sites where the soil fertility is often low. In a glasshouse bioassay, however, all three species not only showed large positive growth responses to supplied P concentrations (0–10mgL ^–1) solutions, but also tolerated high P concentrations (20–40mgL ^–1), well above those normally found in their natural habitat. Plants responded by raising the concentrations of P in their shoot and root tissues and increasing their biomass, resulting in an increased P uptake. A shade-tolerant competitor species, Urtica dioica, also grew vigorously across the full range of P concentrations, restricting the growth of the woodland species. This emphasises the difficulty of establishing semi-natural woodland vegetation in the presence of competitor species, for example in situations where new woodlands are planted on fertile ex-agricultural soils containing large residual concentrations of P. The influence of soil pH on the growth and nutrient relations of A. nemorosa, L. galeobdolon, V. montana, Poa trivialisandU. dioicawas determined in a separate experiment using an ex-arable soil as the growing medium with pH levels adjusted from 7.4 to 5.8 and 4.3 respectively. Acidifying the soil enhanced growth, but reduced the concentrations of N, P and K in the leaves of all three woodland species, probably due to dilution of these minerals in the increased dry matter production. The competitor species (P. trivialis and U. dioica) responded in similar manner to the woodland indicator species. These results suggest that manipulating soil pH as a means of facilitating the establishment of woodland indicator species in new farm woods is unlikely, in the short term, to be effective where competitor species are present.     

Hurricane-induced modification of nitrogen and phosphorus resorption in an aspen clone: an example of diffuse disturbance

Summary On 27 September 1985, Hurricane Gloria intersected the Rhode Island land mass and disrupted normal autumnal element resorption in 20 ramets of a trembling aspen clone (Populus tremuloides). No mechanical injury to the plants was observed, but in contrast to undamaged aspens, the wind/salt-damaged aspens did not withdraw nitrogen (N) or phosphorus (P) from senescing leaves. There was actually 17% more N in abscising leaves than in presenescent, pre-hurricane leaves.