Twenty years of vegetation change in five long-unburned Florida plant communities

Abstract. Rates and directions of change over a 20-yr interval in five long-unburned (> 60 yr) plant communities were studied using multivariate analyses and compositional vectors. The study sites were located in fire and summer-drought adapted, xerophytic vegetation with many endemics on acidic, nutrient-poor, sandy soils in south-central peninsular Florida. Sizes of individual stems from 72 sets of nested permanent quadrats were measured in 1969, 1979, and 1989. Patterns of vegetation change differed by community. Flatwood and bayhead quadrats showed rapid increases in densities and basal areas of Persea borbonia (red bay). In the southern ridge sandhill community, evergreen clonal Quercus species (oaks) and Pinus clausa (sand pine) increased in dominance and grasses declined. Oaks (especially Q. geminata) also increased in importance in scrubby flatwoods. Sand pine scrub was relatively stable in composition, but experienced marked structural changes due to substantial sand pine mortality (18% during 1969–1979, 39% during 1979–1989). Compositional changes in the absence of fire were greatest whereas structural changes were least in southern ridge sandhill and scrubby flatwoods, both communities which normally receive frequent, recurrent fire. Compositional changes were lowest in sand pine scrub, which is normally infrequently burned. Classic successional patterns such as species replacement, decreases in density, and increases in basal area were generally lacking. Tree densities increased in two of four community types (southern ridge sandhill, scrubby flatwoods); while basal area declined in the flatwoods/bayhead and sand pine scrub sites. Directions of compositional vectors included divergent, opposing, and complex patterns, suggesting vegetation change in the absence of fire has a strong stochastic component.     

Damage of alpine vegetation by the 2009 fire on Mt. Shirouma, central Japan: comparison between herbaceous vegetation and Pinus pumila scrub

A fire occurred (0.59 ha) in an alpine fellfield (2600 m a.s.l.) on Mount Shirouma, central Japan, on 9 May 2009 before the start of the growing season. Herbaceous plants and dwarf pine Pinus pumila dominated the site. Plots were established in burned and unburned herb vegetation and P. pumila scrub just after the fire to monitor vegetation recovery. This study reports the short-term monitoring results 3 months after the fire. Burned herb vegetation mostly recovered by late August 2009. However, burned P. pumila did not recover, and other alpine plants were scarce in burned P. pumila scrub. The observed number of species in herb vegetation was 15–20 m⁻² whereas it was only 1–6 m⁻² in P. pumila scrub. The total cover of plants was 111–129% for burned herb vegetation but was only 8–31% for burned P. pumila scrub. Although the species composition in P. pumila scrub distinctly differed between burned and unburned plots, in herb vegetation it was similar between them. Therefore, P. pumila scrub was greatly damaged by the fire, whereas herb vegetation was not damaged. Rapid recovery of herbaceous plants was because winter buds in the soil were not damaged by the fire, but winter buds on shoots of P. pumila were burned. Therefore, the difference in winter bud location (above or belowground) may have resulted in the difference in damage between herbaceous plants and P. pumila.     

Comparison of three devices for estimating fire temperatures in ecological studies

Questions: Are calorimeters and pyrometers accurate and reliable for describing fire parameters in large‐scale ecological projects, and can data from them predict data derived from thermocouples? Do mechanical pre‐treatments in Florida scrub areas alter fire properties? Location: Lake Wales Ridge, Florida, USA. Methods: We deployed thermocouples attached to digital dataloggers, copper and lacquer paint pyrometers and aluminium can calorimeters filled with water in four areas of undisturbed Florida scrub and sandhill vegetation and in adjoining areas pre‐treated with logging, subcanopy felling or mowing. Individual dataloggers were positioned in areas spanning a range of vegetation structure; three calorimeters and three pyrometers were placed with each thermocouple. We also deployed individual calorimeter‐pyrometer pairs. Sites were burned, after which we compared methods for characterizing fire, particularly the ability of pyrometers and calorimeters to predict values derived from thermocouples. Results: Pyrometers best predicted peak 1‐minute mean temperatures and were least successful at estimating maximums. Calorimeters were associated with the number of minutes maximum temperatures exceeded 60 ßC, but were damaged at high residence times and inconsistent at low temperatures. Both pyrometers and calorimeters detected site and treatment differences in fire intensities. These analyses and logistic considerations give pyrometers an edge over calorimeters in situations where dataloggers are impractical. Mechanical pre‐treatments to fire altered fire parameters. Mowing reduced fire temperatures but had different effects on areas burned; this was related to the length of time elapsed between mowing and burning. Subcanopy felling increased both fire coverage and temperatures. Conclusion: Pyrometers outperformed calorimeters as a cheap method for describing relative temperature regimes that are a function of both temperature and residence time. Pyrometers were able to demonstrate how mechanical treatments applied prior to prescribed burning altered fire parameters. Pyrometers are a useful tool for investigating biological responses to fire at multiple scales and in heterogeneous vegetation.     

Gap ecology in Florida scrub: Species occurrence,diversity and gap properties

Questions: Studies of gap effects have been conducted mainly in forests. We studied gap ecology in a pyrogenic Ceratiola ericoides (Florida rosemary) dominated shrubland and asked: How do gap size and the frequency of large gaps change across the fire chronosequence? Do larger gaps differ from smaller gaps in vegetation structure or species diversity? Are effects of gaps independent of, or dependent upon, time‐since‐fire? Are larger gaps refugia for herbs and subshrubs? Location: Archbold Biological Station, Lake Wales Ridge, south‐central Florida, USA. Methods: We investigated plant species occurrence and diversity in 805 gaps (areas free of shrubs taller than 50 cm) in 28 fire‐dependent Florida rosemary scrub sites. We collected quantitative cover data in a subset of seven sites. Results: Gap area distribution was lognormal. The largest gaps occurred throughout all but the longest time‐since‐fire intervals. Gaps were smallest in the longest unburned site but otherwise did not show strong patterns across the fire chronosequence. Species diversity measures increased with increasing gap area, with herbaceous diversity increasing with both gap area and bare sand. Herb diversity (H') decreased with time‐since‐fire. Larger gaps are refugia for some species. Of 14 species occurring in 25–75% of gaps, 13 had increased occupancy with increasing gap area, and gap area was the strongest predictor of occupancy for seven species of herbs and shrubs. Time‐since‐fire was the strongest predictor of occupancy for five species, including four ground lichens that increased with time‐since‐fire. Conclusions: Community structure within Florida scrub gaps is influenced by gap size, which in turn is affected by fire, the dominant ecological disturbance. We present a conceptual model that considers both gap size and time‐since‐fire as drivers of community structure and herbaceous plant diversity in Florida scrub. Because gap properties (independently of fire) have strong influences on species assemblages in Florida rosemary scrub gaps, fire management should consider the number and size of gaps across the landscape.     

Regeneration in fringe mangrove forests damaged by Hurricane Andrew

Mangrove forests along many tropical coastlines are frequently andseverely damaged by hurricanes. The ability of mangrove forests to regeneratefollowing hurricanes has been noted, but changes that occur in vegetationfollowing disturbance by hurricane winds and storm tides have not been studied.We measured changes in plant community structure and environmental variables intwo fringe mangrove forests in south Florida, USA that experienced high windvelocities and storm tides associated with Hurricane Andrew (August1992). Loss of the forest canopy stimulated regeneration via seedlinggrowth and recruitment, as well as resprouting of some trees that survived thehurricane. Initial regeneration differed among species in both forests:Rhizophora mangle L. regenerated primarily via growth ofseedlings present at the time of the hurricane (i.e., release of advancerecruits), but many trees of Avicennia germinans(L.) Stearn and Laguncularia racemosa Gaertn.f.resprouted profusely from dormant epicormic buds. In one forest, which wasformerly dominated by Laguncularia, high densities ofRhizophora seedlings survived the hurricane and grew toform dense stands of saplings and small trees ofRhizophora. In the other forest, there were lowerdensitiesof surviving Rhizophora seedlings (possibly due tohigher storm tide), and extensive bare areas that were colonized byAvicennia, Laguncularia, andherbaceous species. This forest, predominantly Rhizophoraat the time of the hurricane, now contains stands of saplings and small treesofall three species, interspersed with patches dominated by herbaceous plants.These findings indicate that moderately damaged fringe forests may regenerateprimarily via release of Rhizophora advance recruits,leading to single-species stands. In severely damaged forests, seedlingrecruitment may be more important and lead to mixed-species stands.Regeneration of mangrove forests following hurricanes can involve differentpathways produced by complex interactions between resprouting capability,seedling survival, post-hurricane seedling recruitment, and colonizationby herbaceous vegetation. These differences in relative importance ofregeneration pathways, which may result in post-hurricane forestsdifferent from their pre-hurricane structure, suggest that models forregeneration of mangrove forests will be more complex than directregeneration models proposed for other tropical forests whereregeneration after hurricanes is dominated by resprouting.     

Restoration of Florida scrub vegetation in an old field through 23 years after planting

Florida scrub, a fire‐maintained, xeromorphic shrubland, hosts many rare and declining species, but most scrub habitat has been lost to development and agriculture. Formerly cultivated sites offer an opportunity to restore Florida scrub. In this 23‐year study, we test whether scrub species could be restored to an old field and whether scrub vegetation composition and structure could be reestablished over time. Scrub oaks were planted in a section of an old field in 1992 after the site was cleared and treated with herbicide. Additional oaks and other scrub species were planted in 1993. We determined survival and growth annually of a marked sample of scrub species. We sampled vegetation cover in two height strata annually on 10 permanent line‐intercept transects in the old grove and on 20 transects in adjacent, intact scrub. Initial survival of Quercus geminata exceeded that of Q. chapmanii or Q. myrtifolia. Cover of scrub oaks greater than 0.5 m increased from 1.3% in 1992 to 65.2% in 2010. However, the vegetation structure of large Q. geminata did not resemble that of native scrub. Adaptive management led to cutting large oaks in the fall of 2010, and burning the site in early 2011. Cutting and burning appeared to stimulate sprouting and clonal spread of scrub oaks. Ordination analysis indicated directional change related to increasing scrub oak cover and time since planting but the old field still differed from intact scrub. Vegetation has developed toward scrub composition and structure but exotic grasses persist.     

Impacts of Hurricane Frances on Florida scrub-oak ecosystem processes: defoliation, net CO2 exchange and interactions with elevated CO2

Hurricane disturbances have profound impacts on ecosystem structure and function, yet their effects on ecosystem CO₂ exchange have not been reported. In September 2004, our research site on a fire‐regenerated scrub‐oak ecosystem in central Florida was struck by Hurricane Frances with sustained winds of 113 km h⁻¹ and wind gusts as high as 152 km h⁻¹. We quantified the hurricane damage on this ecosystem resulting from defoliation: we measured net ecosystem CO₂ exchange, the damage and recovery of leaf area, and determined whether growth in elevated carbon dioxide concentration in the atmosphere (C_a) altered this disturbance. The hurricane decreased leaf area index (LAI) by 21%, which was equal to 60% of seasonal variation in canopy growth during the previous 3 years, but stem damage was negligible. The reduction in LAI led to a 22% decline in gross primary production (GPP) and a 25% decline in ecosystem respiration (R_e). The compensatory declines in GPP and R_e resulted in no significant change in net ecosystem production (NEP). Refoliation began within a month after the hurricane, although this period was out of phase with the regular foliation period, and recovered 20% of the defoliation loss within 2.5 months. Full recovery of LAI, ecosystem CO₂ assimilation, and ecosystem respiration did not occur until the next growing season. Plants exposed to elevated C_a did not sustain greater damage, nor did they recover faster than plants grown under ambient C_a. Thus, our results indicate that hurricanes capable of causing significant defoliation with negligible damage to stems have negligible effects on NEP under current or future CO₂‐enriched environment.     

Long-term trends in annual reproductive output of the scrub hickory: factors influencing variation in size of nut crop

Reproductive output by the Florida-endemic scrub hickory (Carya floridana Sargent) was studied over a 28-yr period in three south-central Florida vegetation associations: southern ridge sandhill, sand pine scrub, and scrubby flatwoods. The objectives were to describe multi-annual patterns of variation in nut production, identify factors involved in this variation, and investigate differences in patterns among associations. Peaks (higher values bracketed by lower values) in nut production occurred in 7 yr in sandhill and scrub and 8 yr in scrubby flatwoods during the 22-yr period for which we had continuous data. Of the total of 22 peaks in the three associations combined, 17 occurred at intervals of 2 or 3 yr, and peaks occurred in the same years in 18 of the 22 cases. Periodicities of nut production generated by spectral analyses (Fourier transforms) generally agreed with the observed peaks. Numbers of nuts per bearing ramet, proportion of ramets bearing nuts, ramet height, and light availability were positively correlated with nut production. Weather variables, specifically winter rainfall and minimum spring temperatures, accounted for a total of about one-quarter to one-half of the variance in nut production depending on the vegetation association. Following a prescribed fire in a sandhill plot, scrub hickory quickly regained fruit production, but over a 5-yr period following the fire nut production by ramets in the largest size class was reduced compared with the unburned control plot.     

Effects of re‐introducing fire to a central Florida sandhill community

Abstract. A southern ridge sandhill site in central Florida, USA, was burned in 1989, 1991, and 1995 after 63 years of fire‐suppression to simulate a pre‐settlement fire regime. Fire changed species abundance and vegetation structure but caused only minimal changes in species turnover and diversity. There was a general trend for an increase in the cover of herbs following fire but this was a statistically significant effect for only one species, Liatris tenuifolia var. tenuifolia. Aristida beyrichiana increased, litter cover and litter depth were significantly reduced, and ground lichens were eliminated in response to burning. Scrub oaks and palmettos in the ground cover and small shrub layers (height ≤ 1 m) either increased or did not respond to burning, reflecting strong post‐fire resprouting. Diversity in the ground cover and small shrub layers were not affected by fire. Scrub oaks and palmettos in the large shrub and overstorey layers (height > 1 m) were reduced in density, basal area, and longest canopy measurements in response to fire. Species diversity also decreased within these layers following fire. Some Pinus elliottii var. densa survived fire, but their density was reduced. All Pinus clausa were eliminated by fire. Periodic burning can suppress the dominance of shrubs (Quercus spp.) while increasing the cover of grasses and herbs in southern ridge sandhill vegetation.     

Stand structure and dynamics of sand pine differ between the Florida panhandle and peninsula

Size and age structures of stand populations of numerous tree species exhibit uneven or reverse J-distributions that can persist after non-catastrophic disturbance, especially windstorms. Among disjunct populations of conspecific trees, alternative distributions are also possible and may be attributed to more localized variation in disturbance. Regional differences in structure and demography among disjunct populations of sand pine (Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg.) in the Florida panhandle and peninsula may result from variation in hurricane regimes associated with each of these populations. We measured size, age, and growth rates of trees from panhandle and peninsula populations and then compiled size and age class distributions. We also characterized hurricanes in both regions over the past century. Size and age structures of panhandle populations were unevenly distributed and exhibited continuous recruitment; peninsula populations were evenly sized and aged and exhibited only periodic recruitment. Since hurricane regimes were similar between regions, historical fire regimes may have been responsible for regional differences in structure of sand pine populations. We hypothesize that fires were locally nonexistent in coastal panhandle populations, while periodic high intensity fires occurred in peninsula populations over the past century. Such differences in local fire regimes could have resulted in the absence of hurricane effects in the peninsula. Increased intensity of hurricanes in the panhandle and current fire suppression patterns in the peninsula may shift characteristics of sand pine stands in both regions.     

Microhabitat and time-since-fire: effects on demography oF Eryngium cuneifolium (Apiaceae), a Florida scrub endemic plant

Eryngium cuneifolium Small. (Apiaceae) is a narrowly distributed endemic found only in Ceratiola ericoides (Florida rosemary)-dominated Florida scrub, a periodically burned, shrub-dominated habitat. Multivariate analyses using 22 ∗∗∗microhabitat characteristics indicated significant microhabitat and time-since-fire effects on survival, growth, and fecundity of 1287 individuals over a 4-yr period. Survival increased with distance to the nearest shrub, and plants in larger open patches had greater survival rates. Neighboring shrubs of Ceratiola ericoides and Calamintha ashei were associated with a higher mortality of E. cuneifolium than other neighboring shrub species. Survival was reduced by two-thirds over 4 yr (14% vs. 42%) for E. cuneifolium near C. ericoides. Sand accretion increased growth and fecundity. With greater time since fire, woody shrubs increasingly dominate and open patches shrink, significantly reducing survival, growth, and fecundity of E. cuneifolium. Effects were particularly dramatic between 2 and 7 yr postfire, when annual mortality increased from <10% to >30% (r = 0.74). This herbaceous species is dependent on an open habitat maintained by periodic fire. Belowground competition or allelopathy from shrubs probably restricts E. cuneifolium to recently burned, open patches within the most xeric parts of Florida scrub.     

Effects of Hurricane Andrew on stands of slash pine (Pinus elliottii var. densa) in the everglades region of south Florida (USA)

Few hurricanes affect intact stands of subtropical pines. We examined effects of winds in the eyewalls of Hurricane Andrew, where wind speeds were >200 km h^–1, on all remaining large mainland stands of Pinus elliottii var. densa (south Florida slash pine) on limestone outcroppings (rocklands) in the everglades region of southern Florida. We measured densities and sizes of trees and assessed damage and mortality in plots in old-growth stands in the Lostman's Pines (LOP) region of Big Cypress National Preserve and in second-growth stands in the Pines West (PIW) and Long Pine Key (LPK) regions of Everglades National Park. We also examined age-size relationships using sections from trees killed by the hurricane in LOP and LPK. We used the data to predict effects of recurrent hurricanes on the structure and dynamics of the old-growth stand and to compare effects of hurricanes on old- and second-growth stands.Slash pine was resistant to hurricane winds. Most trees in stands (68–76%) were not severely damaged; mortality in the three regions averaged 17–25% shortly after the hurricane and 3–7% during the following year. Mortality was positively associated with tree size; mean tree sizes decreased and size-selective thinning occurred in all stands. Nonetheless, local mortality ranged from 3–4% to 50–60% among plots in all stands. Such local variation in mortality resulted from clustering of large trees, especially in old-growth stands, and from microbursts during the hurricane, which affected all stands. Recurrent, intense hurricanes are predicted to kill larger trees, slowly opening new patches and increasing sizes of extant patches, thus resulting in almost continual presence of openings suitable for recruitment in old-growth stands. Age-size relationships also indicated that large trees in old-growth stands may survive 2–3 centuries. The combination of frequent openings and wind resistance of large trees is predicted to result in old-growth stands that are highly uneven aged, with trees locally distributed in similar-aged patches. The extent to which such stands deviate from demographic equilibrium, as well as turnover rates within stands, are likely to increase as the frequency of recurrent, intense hurricanes increases.Damage and mortality differed in old- and second-growth stands. Large trees were more, but small trees less likely to be damaged in old- than second-growth stands. In contrast, mortality was significantly lower in old- (LOP: 16.9% ± 3.1 [mean ± s.e.]) than second-growth stands (PIW: 22.5% ± 2.0; LPK: 25.2% ± 2.7). Total hurricane-related mortality was 30–60% higher in second- than old-growth stands. Size class structure, more uneven in old- than second growth stands prior to the hurricane, diverged even more afterwards. Hurricane Andrew removed     

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.     

The role of soil seed banks in the restoration of dry acidic dune grassland after burning of Ulex europaeus scrub

Question: Can the seed bank play a significant role in the restoration of plant communities of dry acidic dune grassland where fire has destroyed Ulex europaeus scrub? Location: Northern French Atlantic coast. Methods: One year after the fire, the seed bank and vegetation were sampled in 1 m × 1 m plots along three transects from the oldest scrub vegetation towards the grassland. Differences in species richness, seed density and contribution of ecological groups in the seed bank and vegetation along the transects were analysed. Results: Seed density and species richness in the seed bank decreased significantly from the grassland towards the centre of the scrub vegetation; 50% of the seed bank consisted of core species of the target plant community, such as Carex arenaria, Aira praecox, Rumex acetosella and Agrostis capillaris. Seeds of these species were also found in the deeper soil layers beneath the oldest scrub vegetation, indicating that they can be considered to be long‐term persistent. Beneath the youngest scrub vegetation, seeds of rare satellite target species also occurred. However, no target species were established on the burned site after one year, resulting in a large discrepancy between seed bank and vegetation. Conclusions: Although the seeds present in the soil indicate that restoration of the acidic grassland based on the seed bank is possible, additional management actions such as mowing and soil disturbance may be necessary to restrict resprouting of Ulex and to stimulate the germination of seeds of target species in the deeper soil layers.     

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

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

Demographic effects of fire on two endemic plant species in the longleaf pine-wiregrass ecosystem

Fire can have dramatic effects on the vital rates of plant species and has been used successfully for management in a number of ecosystems. However, the demographic response of species to fire in fire-dependent ecosystems is variable, making it important to study the effects of fire on rare and threatened species. We quantified the effects of fire on Astragalus michauxii and Pyxidanthera brevifolia, two rare endemics of the longleaf pine-wiregrass ecosystem of the southeastern USA, by means of periodic matrix models to project the effect of fire frequency on population growth. In contrast to many species in the longleaf pine-wiregrass ecosystem, fire had short-term negative effects on both species, causing reductions in survival, size, flowering, and fruit production. Relative to the three-year fire intervals to which the study populations are currently exposed, more frequent burning is projected to cause population decline, with the most dramatic effects under annual burning. Although the current longleaf pine-wiregrass ecosystem is fire dependent and has experienced frequent fire for at least several thousand years, we propose that the two endemic species may be remnants from a past vegetation assemblage that experienced less frequent fire and thus may be adapted to longer fire-return intervals compared to other species currently in the ecosystem. Despite the short-term negative effects of fire on the vital rates of these species, longer-term benefits such as reduction of woody encroachment and litter removal may be important for the ultimate success of the species.     

Post-fire seedling establishment in Florida sand pine scrub

Abstract. This study deals with a quantification of pre- and post-fire seedling establishment and microsite characteristics in two Florida sand pine scrub sites burned in May 1993. In addition, life history characteristics related to seedling establishment are described for five perennial species –Calamintha ashei, Chapmannia floridana, Eriogonum floridanum, Garberia heterophylla and Palafoxia feayi. Post-fire seedling establishment in sand pine scrub was sparse (median = 1, 12 seedling/m²), with 17 of 35 species establishing seedlings. Chapmannia, Eriogonum, Garberia and Palafoxia resprouted and flowered after fire; Eriogonum and Garberia had strong post-fire seedling establishment responses within 19 months post-fire. Calamintha individuals were killed by fire, but this species had a strong post-fire seedling establishment response, presumably from seeds in a soil seed bank. Eriogonum and Calamintha seedlings established preferentially in plots centered on conspecific adults. For these species with poor seed dispersal, spatial patterns of seedling establishment may be influenced more by pre-fire adult plant location than by post-fire microsite conditions. Post-fire seedling density in sand pine scrub was much lower than in California chaparral and South African sand plain lowland fynbos.     

Modelling responses of pine savannas to climate change and large‐scale disturbance

Global warming can potentially influence ecological communities through altered disturbance regimes in addition to increased temperatures. We investigate the response of pine savannas in the southeastern United States to global warming using a simple Lotka‐Volterra competition model together with predicted changes to fire and hurricane disturbance regimes with global climate change. In the southeastern United States, decreased frequency of both fires and hurricanes with global warming will shift pine savannas toward a forested state. A CO₂ fertilization effect that increases the growth rate of tree populations will also push southeastern landscapes from open savannas towards closed forests. Transient dynamics associated with climate driven changes in vegetation will last on the order of decades to a century. In our model, the sensitivity of savannas to relative changes in the frequency of fire versus hurricanes is linearly dependent on the growth rate and mortality of trees in fire and hurricane disturbances.     

Spatio‐temporal Analysis of the Effects of Hurricane Ivan on Two Contrasting Epiphytic Orchid Species in Guanahacabibes,Cuba

Hurricanes represent the dominant type of disturbance in many tropical coastal forests. Here, we focus on mortality of epiphytic orchids caused by hurricane Ivan in the Guanahacabibes National Park (Cuba) and subsequent population recovery. We analyzed different aspects of hurricane damage on two contrasting epiphytic orchids, Broughtonia cubensis and Dendrophylax lindenii, as observed in three plots of coastal vegetation and in three plots of semi‐deciduous forest, respectively. First, we quantified the damage to host trees and orchids and explored if hurricane damage depended on height, size, or identity of the host tree. Second, we used mark connection and mark correlation functions to conduct a detailed analysis of small‐scale spatial patterns in hurricane damage for host trees and orchids. Finally, we analyzed the degree of recovery after Ivan during the 6 yr following the storm. Damage of B. cubensis host trees was independent of height and size, but Ivan severely affected larger and higher host trees of D. lindenii. Spatial analysis revealed non‐random structure in damage that differed between species. Broughtonia cubensis exhibited small‐scale spatial correlation in the proportion of damaged orchids, whereas D. lindenii did not. Dendrophylax lindenii showed ‘patchy’ damage patterns, correlated with height, but B. cubensis did not. The relative growth rate of B. cubensis for the 5–17 mo following Ivan was only moderately reduced and fully recovered in subsequent years, whereas that of D. lindenii was severely reduced the first year and did not fully recover thereafter. We hypothesize that differences in the host, vegetation type, and the traits of the two orchids contribute to the different responses to the hurricane.     

Fire-mediated effects of shrubs, lichens and herbs on the demography of Hypericum cumulicola in patchy Florida scrub

Understanding the effect of disturbance and interspecific interactions on population dynamics and availability of suitable habitats for colonization and growth is critical for conservation and management of endangered species. Hypericum cumulicola is a narrowly endemic, small perennial herb virtually restricted to open areas of well-drained white sand in Florida rosemary scrub, a naturally patchy community that burns about every 20–80 years. Over 1 year (September 1994 to September 1995) we evaluated variation in survival, growth and fecundity among 1214 individuals in 14 rosemary scrub patches of different sizes (0.09–1.85 ha) and fire histories (2, 8–10, and >20 years since the last fire). Fire kills aboveground individuals of H. cumulicola, but new individuals were present a year after fire. Recruitment decreased in patches more than a decade post-fire. Survival, annual height growth rate, and fecundity (number of flowers and fruits) were higher in recently burned patches. Scrub patch size did not affect these demographic variables. Survival was positively associated with the presence of conspecifics and negatively related to proximity to the dominant shrub Florida rosemary (Ceratiola ericoides), prior reproductive output, and ground lichen cover. Since H. cumulicola and other herbaceous species in the rosemary scrub depend on sporadic fires to decrease interference of shrubs and ground lichens, its persistence may be threatened by fire suppression. Received: 4 December 1996 / Accepted: 5 June 1997