The role of seed depth,litter, and fire in the seedling establishment of amphicarpic peanutgrass (Amphicarpum purshii)

Summary Amphicarpum purshii is an annual grass which mostly grows in disturbed areas of the New Jersey Pine Barrens, USA. It is amphicarpic, producing spikelets (and seeds) both above and below the soil surface. Previous research has shown that subterranean seed production ensures reproduction in the event of a major disturbance such as fire and results in rapid post-burn colonization of these sandy habitats. The effects of fire, litter, and seed depth were further examined by planting subterranean seeds at four depths in 16 litter-covered flats buried at ground level and comparing plants arising from burned flats with those in undisturbed litter-covered flats. At 0 and 1 cm depth, rates of seedling emergence were lowest in burned flats. Surface-sown seeds produced seedlings more likely to desiccate. Sowing depth had a greater influence on most measured characters than burning treatments. The mean depth of subterranean seed placement by Amphicarpum is 3.5 cm and this coincides with the seed depth from which plants showed the greatest height growth, shoot biomass, and reproductive output. In a second experiment, subterranean seeds on the bare soil surface in clay pots were more likely to lose viability and less likely to germinate than seeds protected by litter or burial in soil. In addition to providing protection from fire, placement of seeds below ground in the sandy habitat of peanutgrass provides conditions more suitable for seed survival and subsequent seedling establishment.     

Interactions between mediterranean shrub species eight years after experimental fire

This paper is part of a wider study in which different combinations of species were assayed for the revegetation of experimentally burnt heathlands. In this particular case, we determined whether there was any interaction between the dominant species, Erica australis, which recovered by vegetative resprouting, and the only successful woody species sown, Cytisus sp., eight years after fire and seven years after sowing. A detailed study was carried out on each square metre of cover and height of both species in three 160 m² plots. Total cover exceeded 100% in all subplots sown with Cytisus, whereas it did not reach 65% in the others, when both species coexist together. There was an inverse correlation between the covers of both species and a positive correlation between the height of Erica australis and the cover and the height of Cytisus. There was no significant difference in Erica cover in plots sown with Cytisus and plots not sown. There could be some competition for light, but both species are capable of coexisting with high canopy overlap, for at least some time. Cytisus is expanding from the seeded subplots, suggesting that the dense cover of Erica australis does not impede the establishment of Cytisus. This revised version was published online in August 2006 with corrections to the Cover Date.     

Competitive effects of the invasive shrub,Lonicera maackii (Rupr.) Herder (Caprifoliaceae), on the growth and survival of native tree seedlings

Invasive plants are often associated with reduced cover of native plants, but rarely has competition between invasives and natives been assessed experimentally. The shrub Lonicera maackii, native to northeastern Asia, has invaded forests and old fields in numerous parts of eastern North America, and is associated with reduced tree seedling density in Ohio forests. A field experiment was conducted to test the effects of established L. maackii on the survival and growth of transplanted native tree species. The experiment examined above-ground competition (by removing L. maackii shoots) and below-ground competition (by trenching around transplanted seedlings). The effects of above-ground competition with L. maackii were generally more important than below-ground competition, though both were detected. Shoot treatment was the key determinant for the survival of all species except P. serotina, whereas trenching only enhanced survival for A. saccharum caged and P. serotina, and only in the shoot removal treatment. For the surviving seedlings, L. maackii shoot removal increased growth of A. saccharum seedlings protected with cages, but actually reduced the growth of unprotected Q. rubra and A. saccharum seedlings, indicating that L. maackii shoots confer some protection from deer browsing. Significant interactions between root and shoot treatment on Q. rubra growth parameters, specifically greatest growth in the shoot present & trenched treatment, is attributed to protection from deer browsing combined with release from below-ground competition. Despite this protective function of L. maackii shoots, the overall effect of this invasive shrub is increased mortality of native tree seedlings, suggesting it impacts the natural regeneration of secondary forests.     

Differential survival of chaparral seedlings during the first summer drought after wildfire

Summary Big Pod Ceanothus (Ceanothus megacarpus) is an obligate seeder after fire; Laurel Sumac (Rhus laurina) is primarily a resprouter after fire. Both species commonly occur together in mixed stands and are dominant members of the coastal chaparral of southern California. We compared the mean survival of post-fire seedlings of each species during the first summer drought after fire and found C. megacarpus to have a mean survival of 54% while R. laurina had a mean survival of only 0.1%. Rooting dephs were similar between species but predawn water potentials and leaf temperatures were higher for R. laurina seedlings. Leaf temperatures for R. laurina reached a mean value of 46.8° C on hot, summer days, about 5° C higher than seedlings of C. megacarpus. By the end of the first growing season, 92% of all C. megacarpus seedlings had suffered herbivory compared to only 17% of all R. laurina seedlings. Herbivory did not appear to be the immediate cause of seedling mortality. Transect data indicated that full recovery of prefire species composition and density at our study site was likely but the mode of recovery was different for the species examined. R. laurina recovered primarily by sprouting, C. megacarpus totally by seedling establishment and a third species, Adenostoma fasciculatum (chamise), by a combination of sprouting and seedling establishment. We attribute the higher mortality of R. laurina seedlings to the greater sensitivity of its tissue to water stress. It may be that differential survival of shrub seedlings and differential modes of reestablishment after fire play an important role in maintaining species diversity in the chaparral communities of coastal, southern California.     

Effects of forest fire ash on germination and early growth of four pinus species

Fire modifies the germination of seeds of numerous species. One of the fire factors that cause these modifications is the ash. This study analysed the germination of seeds of Pinus sylvestris L., Pinus nigra Arn. Pinus radiata D. Don and Pinus pinaster Aiton, subjected to different ash treatments, and the development of seedlings that grew in these conditions for 14 weeks. We obtained the ash by completely burning leaves and small twigs from the most abundant woody species in populations of pines. The ash treatments applied were as follows: Control (without ash), Low (half of the amount registered in a fire), Medium (equal to the amount registered in a fire) and High (double the amount registered in a fire). Germination took place on paper inside Petri dishes and in soil, and growth was only analysed for the plants that grew in the soil. For all species, and especially P. nigra, germination rate decreased as the amount of ash applied increased. The negative effects of the ash were more apparent following treatments in Petri dishes than in the soil. In the dishes, the average germination time varies little between the four species of pine. In soil, the average germination time is more prolonged, and only some differences were observed between treatments in P. sylvestris and P. nigra. In these species the high treatment significantly increased the average germination time. We found a greater sensitivity of P. sylvestris and P. nigra to the addition of ash, showing a higher seedling mortality rate. However, it appears that seedling development (length and weight) was not affected in any of the species through the addition of ash. These tests allow us to deduce that, in P. sylvestris, P. nigra, P. radiata and P. pinaster, the ash produced by forest fires has an inhibiting effect on germination and little effect on the development of seedlings in the first months of life. This revised version was published online in June 2006 with corrections to the Cover Date.     

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

Summary The spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.     

The effects of fire on soil nitrogen associated with patches of the actinorhizal shrub Ceanothus cordulatus

Nitrogen is a limiting resource in many temperate forests and nitrogen-fixing plants are usually limited to the early stages of post-disturbance succession. In fire-dependent Sierra Nevada forests, however, Ceanothus cordulatus is relatively abundant even in old-growth forest conditions which are at least partly maintained by fire. We conducted a field experiment to determine if soil beneath Ceanothus patches represent `resource islands' of available N which persist after fire. Nine plots containing discrete patches of Ceanothus, Arctostaphylos patula (manzanita; chosen as a non N-fixing reference species), and bare forest floor were subjected to either a low-intensity (n=3) or high-intensity (n=3) bum treatment, or remained unburned as controls (n=3). Soil temperatures during the bum were monitored by a network of thermocouples placed at the surface of the mineral soil and at ca. 10 cm depths. Soil samples were collected from the organic horizon, 0-10 cm and 15-25 cm depths within each patch type immediately before burning and 2 days, and 6, and 11 months after. Soil moisture, total C and N, and ammonium and nitrate concentrations were determined in the laboratory. Before the burn, Ceanothus patches were significantly enriched in total and inorganic N in the organic horizon relative to the other patch types. A sharp increase in inorganic N was observed in all patch types and depths immediately following burning, but by 6 months after the burn, Ceanothus patches were significantly enriched relative to the surrounding patch types and remained so at months. Resprouting Ceanothus patches will continue to be an important source of a limiting nutrient in this fire-prone ecosystem.     

Competition effects on wiregrass (Aristida beyrichana) growth and survival

We investigated neighboring plant effects (competition and facilitation)on wiregrass establishment in two frequently encountered restoration situationsof former longleaf pine-wiregrass habitats in the Southeastern United States:longleaf pine plantations and previously cultivated fields. In the plantationexperiment, we specifically examined canopy removal, neighboring wiregrassdensity, and aboveground and belowground effects on establishment and growth ofwiregrass seedlings at two different ages (3 weeks and 6 months) with 3competitive exclusion treatments (aboveground exclusion, belowground exclusion,or no exclusion). Competition treatment effects were age-dependent forsurvivorship and growth. Survival of 6 month old seedlings was unaffected bycompetition treatment; whereas, three week old seedling survival was greatestwhere roots were excluded. Seedling size increased with root exclusion for 6month seedlings, but not for 3 week old seedlings in plots lacking neighboringwiregrass. Where wiregrass was present, both 3 week and 6 month old seedlingsincreased growth with root exclusion. Furthermore, where neighboring wiregrassplants were absent, increasing canopy density resulted in decreased seedlingsize, but did not affect survivorship. In old fields, fertilizer treatments andweeding effects were also assessed using 3 week and 6 month old seedlings.Fertilizer application did not benefit seedling survival or growth and reducedsurvivorship of 3 week old seedlings. Seedlings were smaller where neighboringold field, weedy vegetation was present regardless of wiregrass seedling age;whereas, survival was dependent on seedling age at time of planting. Six monthold seedling survivorship remained high regardless of weeding treatment. It isstill unclear whether fertilizer application is beneficial to the successfulestablishment of wiregrass.     

Effects of leaf litter on woody seedlings in xeric successional communities

We investigated the effect of leaf litter on the establishment of Eucalyptus incrassata, a mallee eucalypt. It has been suggested that litter accumulation may hinder seedling establishment, and that the removal of litter may be one of the mechanisms through which fire enhances recruitment. We conducted factorial experiments testing the effects of three kinds of leaf litter on E. incrassata seeds and seedlings at three contiguous sites with different land use histories. One site was an uncleared E. incrassata open mallee woodland (Mallee site), one a cleared area that had been ungrazed for about five years (Pasture site) and the third an area of mallee rolled some 40 years ago and permitted to regenerate (Regrowth site). Litter had no effect on emergence of planted E. incrassata seeds, but emergence differed between sites. Overall, the percentage of seeds that germinated and emerged was substantial (mean 35.2% ± 25.9%). Seedling shoot biomass did not differ between sites or litter treatments. Although seedlings grown in Pasture litter suffered higher mortality rates, overall mortality rates were low (mean 13.2% ± 15.5%), suggesting that leaf litter has little effect on recruitment rates during winter and spring. We conclude that leaf litter does not affect emergence or growth in young E. incrassata seedlings during winter and spring, when most establishment occurs. Our results emphasize the difficulty in predicting litter effects on recruitment.     

Spatial segregation of pines and oaks under different fire regimes in the Sierra Madre Occidental

Surface fire can modify spatial patterns and self-thinning in pine-oak ecosystems. Spatial pattern analyses were used to compare pattern development and interspecific spatial interactions in trees and seedlings in five Madrean pine-oak stands with different recent fire histories. Interspecific and intraspecific patterns were compared in small (< 15 cm dbh) and large (< 15 cm dbh) diameter classes of the pines (Pinus durangensis, P. teocote, and P. leiophylla) and oaks (Quercus sideroxylla, Q. crassifolia, and Q. laeta) that collectively dominated the five stands. Numbers of juvenile trees in 2.5 × 2.5 m subplots were correlated with cumulative distances to adult trees. Small pine and oak trees were intraspecifically clustered at all scales, irrespective of fire regime. Large pines were strongly clustered only in stands with longer fire-free intervals, and patterns of large versus small pine trees were regular or random in frequent fire stands. These patterns were consistent with fire-induced mortality of maturing trees under frequent fire. Large and small pines were segregated from small oaks at short and long distances in one stand with a 32-year fire-free interval, implying that two or more dynamic factors had produced regular patterns at different scales. Such regular spatial patterns at short distances were not seen in other stands. Therefore, there was little evidence for direct competition between oaks and pines. The results reported here are consistent with studies from other pine-oak ecosystems showing that different fire regime and site factors interact to influence stand development processes and relative dominance of pines and oaks. In some stands, the continued absence of fire could foster increasing tree densities and an intensification of local neighborhood effects, producing segregation of pine and oak species at longer distances. This revised version was published online in August 2006 with corrections to the Cover Date.     

Biotic constraints on the invasion of diffuse knapweed (Centaurea diffusa) in North American grasslands

Knapweeds (Centaurea spp.) are among the most invasive of non-indigenous plant species that have colonized western North America over the last century. We conducted a 4-year experiment in a reconstructed grassland to test hypotheses related to the ability of grasslands to resist the invasion of diffuse knapweed (C. diffusa). We experimentally invaded C. diffusa and three native species into areas where we manipulated soil nitrogen (N) and phosphorus (P) availability and removed extant grasses to reduce competition. We evaluated the growth response of these species to these resources and competitive manipulations. Of the native species that were experimentally added, only one species, Ratibida pinnata (prairie coneflower), established in any numbers. Establishment values in intact vegetation were low for both species, but establishment by C. diffusa (0.02%) clearly outperformed that of R. pinnata (0.001%). Under reduced grass competition, establishment was enhanced, but the values for C. diffusa (0.68%) were not statistically different from those of R. pinnata (0.57%). Neither species performed better under higher soil nutrients in the presence of competing grasses. In plots with both species, biomass of the two planted species was positively correlated, but the biomass of both species was negatively correlated with non-added weedy species. Subsequent harvests of C. diffusa indicated that establishment was enhanced in treatments with higher soil nutrients but that the biomass of these plants could only be enhanced when plant competition was also reduced. These results indicate that C. diffusa can establish in intact grasslands at rates higher than natives, but opportunism rather than competitive ability best describes the invasiveness of C. diffusa. Thus, the mechanisms contributing to the establishment of this knapweed species are different from factors identified as contributing to the dominance of this invader.     

The influence of dominant shrubs,fire, and time since fire on soil seed banks in mixed chaparral

The composition and density of soil seed banks beneath co-occurring Adenostoma fasciculatum and Ceanothus greggii shrubs from three chaparral stands last burned 9, 35 and 85 years before 1986 were investigated. The overall density of seeds in the soil, as estimated by germinations under greenhouse conditions, increased with time since fire (ca. 8000 to 25000/m²). However, this increase was due entirely to the accumulation of A. fasciculatum seed in the soil (ca. 2000 to 21000/m²). In contrast, the density of C. greggii seed was different in each of the three stands, but was not correlated with time since fire: maximum densities were recorded from the 35 year old stand (ca. 2000/m²).A total of 31 taxa germinated and 17 occurred in sufficient numbers to be analyzed statistically. Germinable seed densities of three herb species were not influenced by soil source (beneath A. fasciculatum or C. greggii), time since fire, or the direct effects of a controlled fire treatment. Germinable seed densities of a further nine species were significantly influenced by the elapsed time since stands last burned. The densities of four decreased and five increased. Four of the species that increased in seed density over the three stands were annuals, suggesting that the chaparral sub-canopy habitat is not as unfavorable for annuals as is often assumed. The fire treatment decreased germinable seed densities of four annual species by 40–70%, but increased the germinable seed densities of the shrubs A. fasciculatum and C. greggii, and the annual Phacelia brachyloba. Our results indicate that seeds of A. fasciculatum will increase in the soil bank for at least 85 years after fire in chaparral where it is dominant. In contrast, seed reserves of C. greggii appear to be influenced primarily by site-specific patterns of seed production and by the intensity of post-dispersal seed predation.     

Resprouting vigour of two mediterranean shrub species after experimental fire treatments

Experimental fire treatments were carried out by applying a propane torch flame to individual stumps ofArbutus unedo andErica arborea, two dominant ericaceous shrubs living in relatively moist maquis of the western Mediterranean Basin. No mortality was observed in either species. Individual plant size, measured as the individual stump area, was the most important factor determining both number and biomass of resprouts for all fire treatments. The number of resprouts at 3 months was less at the higher temperature, but no difference could not be detected at 18 months. Duration of flame application had no effect on resprouting success. This effect was not statistically significant 18 months after treatments were applied. Biomass of resprouts was not clearly affected by different fire treatments either 3 months or 18 months after the start of the resprouting process.     

Effects of microsite light availability on the survival and growth of oak seedlings within a grassland

Effects of microsite light availability on the growth and survival of transplantedQuercus serrata Thunb. seedlings in aMiscanthus sinensis Anderss. grass canopy were investigated by a “plant's eye view” approach. Diffuse site factors, i.e., the fractional transmissions of diffuse photosynthetic photon flux density, were estimated at 15 cm aboveground in 108 microsites where the seedlings grew. Microsite diffuse site factors were significantly different between surviving and dead seedlings during the experiment period from April to October (F_[1,14]=10.9, P<0.01). Relative growth rate of dry weight for individual seedlings positively depended on the diffuse site factors (r²=0.482, P<0.001 in May; r²=0.312, P<0.001 in October). Only 16 seedlings produced their second stem flush within the grass canopy. The ratio of height to dry weight of the second stem flush was significantly higher for the seedlings grew in shady microsites than for those in less shady microsites (r²=0,471, P<0.01 in May). This study suggests that the microsite heterogeneity of light availability is one of the important factors affecting the establishment of tree seedlings in patchy grasslands.     

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.     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

Small-scale spatial heterogeneity in the vertical distribution of soil nutrients has limited effects on the growth and development of Prosopis glandulosa seedlings

We examined the above- and below-ground responses of seedlings of the woody shrub Prosopis glandulosa to the spatial heterogeneity of soil nutrients within the root zone. We performed a microcosm experiment where seedlings were grown with different combinations of nutrients [nitrogen (N), phosphorus (P), and both combined (NP)] and under different levels of nutrient heterogeneity (nutrients supplied as patches located in the bottom and/or upper portion of rooting zone versus homogeneous distribution). Seedling morphology and biomass did not show a strong response to changes in nutrient ion or spatial heterogeneity. Height, number of leaves, and specific leaf area did not vary significantly between treatments. The number of leaves, foliar biomass, stem biomass and biomass allocation to stems of seedlings showed more responsiveness to the addition of N and NP than to the addition of P. The spatial heterogeneity of nutrients affected the diameter, root biomass and leaf N content. Seedlings had higher diameter and root biomass when the nutrients were homogeneously distributed as compared to their placement as patches in the bottom of the microcosms. Their leaf N concentration increased in those treatments where the nutrient patch was located in the lower half as compared to the upper half of the microcosms. Root foraging responses to nutrient patches varied with their location. Significant root proliferation was observed when patches of N, P and NP were located in the upper portion of the rooting zone; when they were located in the lower portion such a response was observed only for P. Despite our findings that Prosopis seedlings have a low overall responsiveness to small-scale vertical differences in soil nutrient heterogeneity, our results suggest that these differences may modify the growth dynamics of the secondary roots of this ecologically important dryland species during the early stages of its development.     

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.     

Factors controlling postfire seedling establishment in southern California chaparral

Summary This paper discusses the interactions among prefire shrub abundance, soil moisture, and plant and animal species on postfire seedling establishment in mixed chparral in southern California. Postfire germination and survival of seedlings in a stand dominated by a facultative seeder (Adenostoma fasciculatum) and by an obligate seeder (Ceanothus greggii) were monitored for 2 years. Relative to prefire abundance, germination of C. greggii was higher than that of A. fasciculatum. Survival during the first year was also higher in C. greggii than in A. fasciculatum. During the second year, however, mortality of C. greggii was greater than that of A. fasciculatum, mostly due to a psyllid infection. Germination of A. fasciculatum was negatively related to prefire shrub abundance. C. greggii germination was not associated with prefire shrub abundance. Seedling mortality of both species was very strongly related to the depletion of soil moisture the first few months after germination. A. fasciculatum was more sensitive than C. greggii to the drying of the soil, especially in the upper levels. C. greggii seedlings had longer roots, greater root/shoot biomass ratios, higher water potentials, and a later peak in seasonal growth activity compared to A. fasciculatum. Herbs promoted greater survival of A. fasciculatum. Our results indicate that the obligate seeder species, C. greggii, is better adapted to establish seedling in chaparral by producing greater relative germination and greater seedling survival than the facultative seeder species A. fasciculatum. The greater adaptability of C. greggii to the physical environment is counteracted by interspecific interference by plant and animal interactions which tend to favor A. fasciculatum over C. greggii.     

Survivorship of penstemon grandiflorus in an oak woodland: combined effects of fire and pocket gophers

Summary The effects of fire and pocket gophers, Geomys bursarius, on the survivorship of Penstemon grandiflorus growing in an oak woodland in Minnesota were studied from 1986 to 1990. Plants growing in sparse vegetation experienced mortality rates twice that of plants growing in dense vegetation. This difference was due partly to pocket gophers whose earth moving activities reduce the density of vegetation and bury and kill individual Penstemon plants. Laboratory feeding trials showed that gophers readily eat Penstemon, particularly the fleshy roots. An experiment involving the removal of 25–75% of the root tissue in 90 plants showed that root loss significantly reduced survivorship, suggesting that gopher herbivory might also kill plants. When gophers were experimentally excluded, plants growing in sparse vegetation exhibited significantly lower mortality rates than those growing in dense vegetation. Plants in the smallest size class exhibited reduced survivorship following a late spring burn; however, overall patterns of survivorship of plants in burned areas did not differ markedly from those in the unburned areas. A longitudenal analysis of plants with different reproductive histories revealed no survivorship cost to reproduction. Mortality rates decreased with increasing plant size. Small plants were more likely to be killed by fire and by being buried under gopher mounds. Differences in underground energy reserves of small and large plants can account for most of the survivorship patterns observed in this study. The study shows that within openings of the oak woodland, fire and gophers reduce the survival of individual Penstemon plants. Nevertheless, since both gophers and fire also serve to perpetuate suitable habitat in the woodland, Penstemon is ultimately dependent on both for its long term persistence in the landscape.