Recovery patterns of three chaparral shrub species after wildfire

Summary In a mature, even aged stand of mixed chaparral, Rhus laurina (facultative resprouter) had consistently higher water potentials and deeper roots than Ceanothus spinosus (facultative resprouter) and Ceanothus megacarpus (obligate seeder). For two years following a wildfire, the same stand of chaparral had resprouts with higher survivorships, predawn water potentials, stomatal conductances, photosynthetic rates and shoot elongation rates than seedlings. Supplemental irrigation of seedlings during summer months removed differences between resprouts and seedlings suggesting that the cause of such differences was limited water availability to the shoot tissues of seedlings. After two years of postfire regrowth, mean seedling survivorship for the obligate seeder (C. megacarpus) was 42%, whereas seedling survivorship for facultative resprouters was only 18% (C. spinosus) and 0.01% (R. laurina). Our results are consistent with the hypothesis that lack of resprouting ability among obligate seeders is offset by an enhanced ability to establish seedlings after wildfire, allowing obligate seeders to maintain themselves in mixed populations through many fire cycles.     

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.     

Species-mediated soil moisture availability and patchy establishment of Pseudotsuga menziesii in chaparral

The occurrence of mature individuals of Pseudotsuga menziesii in stands of Arctostaphylos species mark the initial stages of mixed evergreen forest invasion into chaparral in central coastal California. We planted two cohorts of P. menziesii seeds at three sites under stands of two Arctostaphylos species and Adenostoma fasciculatum in order to determine whether first-year seedling emergence and survival, particularly during the regular summer drought, underlie the spatial distribution of mature trees observed in chaparral. Regardless of the chaparral species they were planted under, P. menziesii seeds that were not protected from vertebrate predation displayed very little emergence and no survival. In contrast, emergence of P. menziesii that were protected from vertebrate predators was much higher but still did not significantly differ among the three chaparral species. However, survival of protected seedlings under Arctostaphylos glandulosa was much greater than under A. fasciculatum, with intermediate survival under Arctostaphylos montana. While mortality of protected seedlings due to insect herbivory, fungal infection, and disturbance displayed no consistent patterns, summer drought mortality appeared to drive the patterns of survival of P. menziesii under the different chaparral species. These emergence, mortality, and survival data suggest that spatial patterns of P. menziesii recruitment in chaparral are driven by first-year summer drought seedling mortality, but only in years when seeds and seedlings are released from vertebrate predation pressure. Because the first-year drought mortality and survival patterns of P. menziesii seedlings differed strongly depending on the chaparral species, we examined the additional hypothesis that these patterns are associated with differences in the availability of soil moisture under different chaparral species. Both higher survival and lower drought mortality of P. menziesii seedlings were associated with higher soil water potential under Arctostaphylos stands during the summer drought, especially in the subsurface soil. The data suggest that Arctostaphylos stands, particularly stands of A. glandulosa, ameliorate xeric summer conditions to a degree that facilitates first-year establishment of P. menziesii and strongly influences spatial distribution of mature trees. Received: 18 September 1998 / Accepted: 23 December 1998     

Tissue water relations of three chaparral shrub species after wildfire

Summary We compared the tissue water relations among resprouts and seedlings of three chaparral species during the first summer drought after wildfire. Two of the species, Rhus laurina and Ceanothus spinosus recover after fire by a combination of resprouting and seedling establishment (facultative resprouters), whereas a third species, Ceanothus megacarpus recovers by seedling establishment alone (obligate seeder). Our objectives were to document any differences in tissue water characteristics that might arise between resprouts and seedlings and to test the hypothesis that seedlings of obligate seeders develop more drought tolerant characteristics of their tissues than seedlings of facultative resprouters. We found that resprouts had much higher predawn values of water potential, osmotic potential, and turgor potentials than seedlings. Predawn turgor potentials of resprouts were 1.5 MPa through July and August when turgor potentials for seedlings remained near 0 MPa. During summer months, midday water potentials were 2 to 3 MPa higher for resprouts than seedlings and midday conductances of resprouts were two to five fold greater than those of seedlings. Even though resprouts did not experience severe water stress like seedlings, their tissue water characteristics, as determined by pressure-volume curve analyses, were similar by the peak of the drought in August. Further-more, the tissue water characteristics of seedlings from the obligate seeder, C. megacarpus, were similar to those of facultative resprouters — R. laurina, and C. spinosus. We attribute the observed differences in plant water status between resprouts and seedlings to differences in rooting depths and access to soil moisture reserves during summer drought. We conclude that the higher growth rates, photosynthetic performance, and survivorship of postfire resprouts are primarily a result of higher water availability to resprouting tissues during summer months. It appears that the greater seedling survivorship during summer drought observed for the obligate seeder, C. megacarpus, is not associated with more favorable tissue water characteristics.     

POST-FIRE REGENERATION OF SOUTHERN CALIFORNIA CHAPARRAL

Succession of chaparral shrubs was studied for the first 4 years after fire on the four major slope faces at three elevations in southern California. Although total cover fluctuated from year to year, shrub cover increased annually through the third year. There was little or no increase in shrub cover between the third and fourth years. Four years after fire the shrubs covered 55% ground surface at the highest elevation site but only 28% at the lowest elevation site. Shrub cover was similar between slope faces with one exception; at the lowest elevation site shrub cover was twice as great on the north and east-facing slopes as on the south and west-facing slopes. Post-fire recovery of shrubs was by seedlings and/or resprouts. Yucca whipplei was an exception in that it did not resprout once the aboveground parts were killed nor did seedlings establish after fire, however the aboveground parts of many Y. whipplei survived the fire. Species which reproduced entirely by seed did so in the first post-fire year from soil-stored seed with the exception of Ceanothus greggii at the highest elevation site. This species was entirely absent the first year after fire but abundant in the second year. Species producing both resprouts and seedlings varied from site to site in the proportion of resprouts:seedlings. Between 83–100% of the post-fire populations of Cercocarpus betuloides, Arctostaphylos glandulosa, and Xylococcus bicolor were resprouts whereas 12–13% of Ceanothus tomentosus were resprouts. For Adenostoma fasciculatum, resprouts constituted 27–54% of the population at the lowest elevation and 65–94% at the highest elevation; whereas, the Quercus dumosa population was 100% resprouts at the lowest elevation and 31–67% resprouts at the highest elevation. Data are presented on the height of resprouts and seedlings for all species at the end of the first post-fire year.     

Post-fire regeneration strategies of Californian coastal sage shrubs

Summary Regeneration methods for coastal sage srub vegetation after fire were studied in the coastal Santa Monica Mountains of southern California. Six sites were sampled two years after a large fire of fall, 1978. The intensity of fire varied. Foliar cover and flowering incidence were recorded for individuals regenerating by resprouting or from seed. Resprouting plants contributed most to post-fire recovery, comprising 95% of the relative foliar shrub cover; 84% of resprout and 47% of seedling cover had flowered. An ANOVA of reproductive mode and fire intensity indicates that resprout total cover and individual size are significantly greater than those of seedlings, regardless of fire intensity. Among sites the average foliar cover of resprouts exceeded that of seedlings by factors ranging from 9 to 63. All coastal sage species examined resprout, although the potential vigor of resprouting appears to vary widely within genera (e.g. Encelia, Eriogonum, and Salvia) and even within species. In the second growing season following fire seedling density increased due to seeds shed by resprouted shrubs. Most of the cover on these stands of coastal sage scrub is destined to be either crown-sprouted individuals or their progeny.     

An individual-based model of chaparral vegetation response to frequent wildfires

The Santa Monica Mountains are home to many species of chaparral shrubs that provide vegetative cover and whose deep roots contribute to the stability of the steep slopes. Recently, native chaparral have been threatened by an unprecedented drought and frequent wildfires in Southern California. Besides the damage from the wildfires themselves, there is the potential for subsequent structural losses due to erosion and landslides. In this paper, we develop a mathematical model that predicts the impact of drought and frequent wildfires on chaparral plant community structure. We begin by classifying chaparral into two life history types based on their response to wildfires. Nonsprouters are completely killed by a fire, but their seeds germinate in response to fire cues. Facultative sprouters survive by resprouting but also rely on seed germination for post-fire recovery. The individual-based model presented here simulates the growth, seed dispersal, and resprouting behavior of individual shrubs across two life history types as they compete for space and resources in a rectangular domain. The model also incorporates varying annual rainfall and fire frequency as well as the competition between plants for scarce resources. The parameters were fit using seedling and resprout survivorship data as well as point quarter sampling data from 1986 to 2014 at a biological preserve within the natural landscape of the Malibu campus of Pepperdine University. The simulations from our model reproduce the change in plant community structure at our study site which includes the local extinction of the nonsprouter Ceanothus megacarpus due to shortened fire return intervals. Our simulations predict that a combination of extreme drought and frequent wildfires will drastically reduce the overall density of chaparral, increasing the likelihood of invasion by highly flammable exotic grasses. The simulations further predict that the majority of surviving shrubs will be facultative sprouting species such as Malosma laurina.     

Regeneration of a dominant Mediterranean dwarf-shrub after fire

Abstract. The regeneration after fire of the dominant east Mediterranean dwarf-shrub Sarcopoterium spinosum was studied under various habitat and geographic conditions in northern Israel, northern Crete and the Aegean coast of Turkey. Age structures of regenerating stands were determined 3–9 yr after fire on 13 1 mx 1 m sample stands by analysis of growth rings in the taproot crown of 874 plants. S. spinosum regenerated either by massive seedling recruitment in some habitats or by resprouting from surviving meristems in others. This versatility facilitates adaptation of the species to widely different environmental conditions and is a factor in the maintenance of its dominance after fire at occupied sites. In the years after fire, the growth of younger plants within the stand at all sites was severely suppressed by the older shrubs. Consequently, the age structure of the stands was dominated by one or two older cohorts that became established soon after fire. In addition, on most habitats, older shrubs (usually older than 6 yr) were often composed of clusters of plants that suggested clonal proliferation of ramets. The strongly peaked age structure, together with the appearance of clonal regeneration on the older plants, suggests that subsequent maintenance of the stand was not dependent on recurrent seedling recruitment. This can explain the persistence of S. spinosum in stands of herbaceous vegetation despite the extreme sensitivity of the seedlings to competition under such conditions.     

Water use patterns of four co-occurring chaparral shrubs

Summary Mixed stands of chaparral in California usually contain several species of shrubs growing close to each other so that aerial branches and subterranean roots overlap. There is some evidence that roots are stratified relative to depth. It may be that root stratification promotes sharing of soil moisture resources. We examined this possibility by comparing seasonal water use patterns in a mixed stand of chaparral dominated by four species of shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a neutron probe and soil phychrometers to follow seasonal depletion and recharging of soil moisture and compared these patterns to seasonal patterns of predawn water potentials, diurnal leaf conductances, and diurnal leaf water potentials. Our results indicated that 1) Quercus was deeply rooted, having high water potentials and high leaf conductances throughout the summer drought period, 2) Heteromeles/Adenostoma were intermediate in rooting depth, water potentials, and leaf conductances, and 3) Rhamnus was shallow rooted, having the lowest water potentials and leaf conductances. During the peak of the drought, predawn water potentials for Quercus corresponded to soil water potentials at or below a depth of 2 m, predawn water potentials of Heteromeles/ Adenostoma corresponded to a depth of 0.75 m, and predawn water potentials of Rhamnus corresponded to a depth of 0.5 m. This study supports the concept that co-occurring shrubs of chaparral in California utilize a different base of soil moisture resources.     

A Population Model of Chaparral Vegetation Response to Frequent Wildfires

The recent increase in wildfire frequency in the Santa Monica Mountains (SMM) may substantially impact plant community structure. Species of Chaparral shrubs represent the dominant vegetation type in the SMM. These species can be divided into three life history types according to their response to wildfires. Nonsprouting species are completely killed by fire and reproduce by seeds that germinate in response to a fire cue, obligate sprouting species survive by resprouting from dormant buds in a root crown because their seeds are destroyed by fire, and facultative sprouting species recover after fire both by seeds and resprouts. Based on these assumptions, we developed a set of nonlinear difference equations to model each life history type. These models can be used to predict species survivorship under varying fire return intervals. For example, frequent fires can lead to localized extinction of nonsprouting species such as Ceanothus megacarpus while several facultative sprouting species such as Ceanothus spinosus and Malosma (Rhus) laurina will persist as documented by a longitudinal study in a biological preserve in the SMM. We estimated appropriate parameter values for several chaparral species using 25 years of data and explored parameter relationships that lead to equilibrium populations. We conclude by looking at the survival strategies of these three species of chaparral shrubs under varying fire return intervals and predict changes in plant community structure under fire intervals of short return. In particular, our model predicts that an average fire return interval of greater than 12 years is required for 50 % of the initial Ceanothus megacarpus population and 25 % of the initial Ceanothus spinosus population to survive. In contrast, we predict that the Malosma laurina population will have 90 % survivorship for an average fire return interval of at least 6 years.     

Causes of arrested succession in coastal dune forest

Dispersal or recruitment limitation may arrest succession after disturbance. In north-eastern South Africa the Acacia karroo successional pathway is used to facilitate coastal forest recovery after strip-mining. However, although A. karroo establishes naturally, it forms monospecific stands, arresting forest succession for decades. This casts doubt on the efficacy of this restoration pathway. We investigated the causes of arrested succession. The seed and seedling banks of A. karroo stands and of forest at Cape Vidal, and three A. karroo stands (7–27 years old) on rehabilitated strip-mined dunes at nearby Richards Bay were examined. The establishment and growth of seedlings at Cape Vidal were also considered. The seed bank was larger and more diverse in forest, but the seedling bank was larger in Acacia stands. At Richards Bay, the size of the seed bank increased and the seedling bank decreased with Acacia stand age. Excluding mammalian herbivores in Acacia stands at Cape Vidal resulted in greater species richness and survival of naturally established seedlings, as well as two experimentally planted species. Neither seed dispersal nor seedling establishment limited recruitment of tree species in Acacia stands. Herbivory arrested forest succession by causing the differential mortality of seedlings. In contrast, at Richards Bay where there were few mammalian herbivores, the advanced regeneration in A. karroo stands converged on the diversity of nearby forests 29 years after restoration. Controlling herbivore access and seeding Acacia stands with forest species are site-specific options for preventing arrested succession when using the A. karroo successional pathway.     

Change over 70 years in a southern California chaparral community related to fire history

Question: What changes in species composition and cover have occurred in chaparral as a function of fire history across an ecoregion? Location: San Diego County, California, USA. Methods: Stands in which 40 mid‐elevation chaparral vegetation plots (each 400 m² in area) were located in the 1930s were resurveyed in 2001. We stratified the stands into Infrequently versus Frequently burned (0–1 versus 2 or more fires recorded in the 91‐yr period), and Immature versus Mature (ã31 yr versus >31 yr since last fire), resulting in four groups. Ten stands were randomly selected from each of these groups for survey. Results: There were no major shifts in life form composition, e.g., live oak trees were not invading chaparral that had experienced little or no fire, nor were subshrubs or herbaceous species replacing shrubs in areas that had experienced more frequent fires. However, there was a notable increase in the frequency of the subshrub Eriogonum fasciculatum across all fire history groups. In the mature stands with infrequent fire, average cover of resprouting shrubs increased (from 72 to 91%) and cover of obligate seeding shrubs (species with fire‐cued germination) decreased (from 21 to 6%) significantly. Mature stands with frequent fire showed a significant decrease in resprouter cover (from 87 to 80%) and increase in obligate seeders (from 10 to 16%). Conclusions: While the tremendous changes in land use in southern California have been predicted to cause shifts in chaparral composition, these shifts are difficult to detect because species longevity and fire cycles are on the order of decades to a century. In this study, the expected trends could only be detected in groups that were mature at the time of the second survey.     

Population structure of Adenostoma fasciculatum in mature stands of chamise chaparral in the southern Sierra Nevada,California

Summary In the low elevation chaparral areas of Sequoia National Park, California, pure stands of chamise (Adenostoma fasciculatum) are periodically rejuvenated by fire. Mature stands showed considerable variability in density and total biomass even though a positive correlation exists between the two. Mature stands showed a preponderance of individuals in the smaller size classes (inverse-J shape distribution). Dead shrubs found in mature stands also tended to be in the smaller size classes. This relatively high mortality of small individuals is important to post-fire stand development. In addition, resprout and seedling biomass one year after fire both showed inverse-J shaped size-class structures. A positive correlation existed between the preburn basal area of a shrub and its first year resprout biomass. Shrub biomass and distance to nearest neighbor were poorly correlated. A significant correlation existed between stand density and a stand's variance-to-mean ratio, indicating a trend toward more regular spacing as density increases. Pre-burn and fire-induced mortality tended to move the stand towards a more clumped distribution. Seedlings replaced dead individuals after a fire and thus restored regular spacing.     

Nitrogen fixation by Cercocarpus ledifolius (Rosaceae) in pioneer habitats

Summary Soil properties of pioneer Pinus flexilis stands with similar topography and climate were investigated. Soils supporting this tree in association with Cercocarpus ledifolius were found to have higher percentages of total nitrogen than soils beneath similar stands lacking Cercocarpus.An excavated Cercocarpus ledifolius shrub in a Pinus flexilis stand in the San Bernardino Mountains of California was found to be nodulated and these nodules were found to be capable of fixing nitrogen. Other known nitrogen-fixing shrubs are frequent associates of Pinus flexilis in extreme sites.Implications of the phylogenetic relationship of some nitrogen-fixing species are briefly discussed, as well as their frequent occurrence in pioneer or extreme habitats.     

Restoring Quercus pyrenaica forests using pioneer shrubs as nurse plants

Question: How to improve reforestation success of Quercus pyrenaica. Location: 1800 m a.s.L, southern Spain. Methods: One‐year‐old Quercus pyrenaica seedlings were planted using two treatments: (1) bare soil, using a 30‐cm diameter augur bit (conventional technique) and (2) under the canopy of a pioneer shrub, Salvia lavandulifolia, using a 12‐cm diameter augur bit. Survival and growth were monitored for six years. Our hypothesis is that the use of shrubs as nurse plants is an alternative technique of reforestation with higher success than traditional techniques, in which pre‐existing vegetation is usually considered a source of competition. The rationale for the study was that for environments with a dry season, pre‐existing vegetation buffers summer drought stress, ameliorates the water status of seedlings and thus usually increases seedling recruitment. Results: Quercus survival was 6.3 × higher when planted under individuals of the pioneer shrub as compared to open areas. Quercus seedlings under shrubs also had shoots 1.8 X longer, while the number of shoots per plant did not differ among treatments. The first summer was the period with the highest mortality (49.1% of seedlings). Summer drought was the main cause of mortality. Conclusions: The use of shrubs as nurse plants for Q.pyrenaica reforestation is a viable technique to increase establishment success. The technique could be similarly useful in other environments with a dry period and for other Quercus species. In addition, this technique offers the advantage of following natural succession, thus minimizing the impact in the community.     

Canopy vs. soil effects of shrubs facilitating tree seedlings in Mediterranean montane ecosystems

Abstract. Question: Is the facilitative effect of nurse shrubs on early recruitment of trees mediated by a ‘canopy effect’(microclimate amelioration and protection from herbivores), a ‘soil effect’(modification of soil properties), or both? Location: Two successional montane shrublands at the Sierra Nevada Protected Area, SE Spain. Method: Seedlings of Quercus and Pinus species were planted in four experimental treatments: (1) under shrubs; (2) in open interspaces without vegetation; (3) under shrubs where the canopies were removed; (4) in open interspaces but covering seedlings with branches, mimicking a shrub canopy. Results: Both effects benefited seedling performance. However, microclimatic amelioration due to canopy shading had the strongest effect, which was particularly pronounced in the drier site. Below‐ground, shrubs did not modify soil physical characteristics, organic matter, total N and P, or water content, but significantly increased available K, which has been shown to improve seedling water‐use efficiency under drought conditions. Conclusions: We propose that in Mediterranean montane ecosystems, characterised by a severe summer drought, pioneer shrubs represent a major safe site for tree early recruitment during secondary succession, improving seedling survival during summer by the modification of both the above‐and below‐ground environment.     

Structural determinants of increased susceptibility to dehydration‐induced cavitation in post‐fire resprouting chaparral shrubs

It is well established that transpiration and photosynthetic rates generally increase in resprouting shoots after fire in chaparral shrublands. By contrast, little is known about how plant hydraulic function varies during this same recovery period. We hypothesized that vascular traits, both functional and structural, would also shift in order to support this heightened level of gas exchange and growth. We examined stem xylem‐specific hydraulic conductivity (K_s) and resistance to cavitation (P₅₀) for eight chaparral shrub species as well as several potential xylem structural determinants of hydraulic function and compared established unburned plants and co‐occurring post‐fire resprouting plants. Unburned plants were generally more resistant to cavitation than resprouting plants, but the two groups did not differ in K_s. Resprouting plants had altered vessel structure compared with unburned plants, with resprouting plants having both wider diameter vessels and higher inter‐vessel pit density. For biomechanics, unburned plants had both stronger and denser stem xylem tissue than resprouting plants. Shifts in hydraulic structure and function resulted in resprouting plants being more vulnerable to dehydration. The interaction between time since disturbance (i.e. resprouting versus established stands) and drought may complicate attempts to predict mortality risk of resprouting plants.     

Fire intensity and herbivory effects on postfire resprouting of Adenostoma fasciculatum in southern California chaparral

Summary Resprouting is the main regeneration mechanism after fire in Mediterranean-type ecosystems. Herbivores play an important role in controlling postfire seedling establishment, but their influence on regeneration by resprouting is less well known. To study the effects of fire intensity on resprouting of Adenostoma fasciculatum in southern California chaparral, and its interaction with herbivory, we conducted an experimental burn at three levels of fire intensity. We found that increasing fire intensity increased plant mortality, reduced the number of resprouts per plant, and delayed the time of resprouting. Herbivory increased with fire intensity, and was related to the time of resprouting. Plants resprouting later in the season and out of synchrony with the main flush were attacked more readily by herbivores. Post-resprouting mortality also increased with fire intensity and was significantly associated with herbivory in the higher fire intensity plots. Fire intensity effects on chaparral regeneration by resprouting may be farreaching through effects on the population structure, resprout production, and growth of Adenostoma fasciculatum.     

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.     

Posthurricane Seedling Structure in a Multi‐aged Tropical Dry Forest: Implications for Community Succession

Hurricane‐caused tree mortality in tropical dry forests occurs predominantly in early successional species. Consequently, hurricanes may accelerate succession in these forests. Forest regeneration, however, must be measured over an extended posthurricane time period to demonstrate this pattern. In this study, we recorded tree seedlings in 19 Florida Keys forests during May–August 1995, 3 years after Hurricane Andrew. For these forests—spanning a chronosequence from 14 to over 100 years since the most recent clearing—we used weighted averaging regression on relative abundances of pre‐hurricane trees to calculate a successional age optimum for each species; and used weighted averaging calibration to calculate inferred successional ages for stands based on pre‐hurricane trees and on posthurricane seedlings. To test the hypothesis that successional stage of seedlings exceeded successional stage of pre‐hurricane trees, we compared inferred stand ages based on posthurricane seedlings with those based on pre‐hurricane trees. Across the study area, inferred stand ages based on posthurricane seedlings were greater than those based on pre‐hurricane trees (P < 0.005); however, more seedlings in the youngest stands were early successional than in older stands. Of 29 species present both as pre‐hurricane trees and posthurricane seedlings, 23 had animal‐dispersed seeds. These results provide evidence that: (1) hurricanes do not ‘reset’ succession, and may accelerate succession; and (2) a strong legacy of stand successional age influences seedling assemblages in these forests.