Effects of repeated burning on species richness in a Florida pine savanna: A test of the intermediate disturbance hypothesis

Abstract. We studied the effect of burning frequency on the density and species richness of understory flowering stems in a Florida sandhill. Flowering stems were censused weekly for 54 weeks in six sites that had been burned one to six times in the previous 16 years. We concurrently measured overstory characteristics such as species composition, density and basal area. We used maximum likelihood and Akaike's Information Criterion to compare linear, quadratic, saturating, and null models of community response to repeating burning. We did not find a relationship between species richness, diversity or flowering stem density and fire frequency. Tree density was related to fire frequency and may represent an indirect pathway for fire effects on understory characteristics. While we found no support for the Intermediate Disturbance Hypothesis, an analysis of our experimental design indicated that we had low statistical power. We develop the hypothesis that a saturating model of response to fire best describes understory species richness in our system. We test this hypothesis using the most extensive published fire data set we are aware of and find support for a saturating model.     

Facilitation of tree seedling establishment in a sand dune succession

Abstract. A nucleated pattern of establishment by pine seedlings of Pinus strobus and P. resinosa around pre-established oak trees of Quercus rubra in a sand dune succession in Ontario, Canada was examined using field observations, seed planting and habitat manipulation. Densities of young pines beneath oak canopies were approximately six times greater than in treeless areas, and densities on the north sides of the trees were significantly greater than on the south sides. However, oaks younger than 35 yr showed no preferential establishment beneath them, while the pine population structures beneath older oaks indicated single periods of successful recruitment. Pine seed planted beneath and beyond oak canopies of three sizes germinated primarily beneath the canopies of medium and large-sized oaks, but subsequent survivorship of seedlings over two growing seasons was poor. Several micro-environmental conditions were changed by oak canopies, but only shade showed a pattern closely corresponding to that of seedling establishment. A habitat manipulation experiment confirmed the primary role of shade in facilitating pine seedling establishment beneath oaks. Failure of pine to continue recruiting successfully beneath facilitating trees is tentatively attributed to intraspecific competition among pine individuals.     

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.     

Post-fire bryophyte dynamics in Mediterranean vegetation

Abstract. Bryophyte dynamics after fire in the Mediterranean macchia of Southern Italy was studied both by diachronic and synchronic approaches. Changes of bryophyte cover and species composition were found in relation to both age and fire intensity. During the first 2 yr after fire, bryophytes dominated the plots which had experienced the highest fire intensity while herbs were dominant in plots affected by lighter fires. Pioneer species, such as Funaria hygrometrica, Barbula convoluta and Bryum dunense, characterized recent intense fires, whereas Bryum torquescens, B. radiculosum and B. ruderale were dominant after less intense burning. Pleurochaete squarrosa, Tortula ruraliformis and Tortella flavovirens dominated intermediate successional stages. Pleurocarpous mosses were dominant only in the older closed stands. Different patterns of regeneration strategies were described: spores dominated early stages of intense fire, while vegetative propagules characterized later successional stages and less severely burned areas. Although bryophytes usually have a low abundance in Mediterranean vegetation, their role in post-fire vegetation dynamics may be locally enhanced according to burning conditions.     

Microsite variation in light availability and seedling growth ofQuercus serrata in a temperate pine forest

Spatial heterogeneity in light availability for tree seedlings under the canopy of a temperate pine forest was studied. Six-day measurements at 10-s intervals revealed a great variety in the temporal patterns of photosynthetic photon flux density (PPFD) and histograms among observation days and microsites; mean daily total PPFD relative to full sun varied from 1.5% to 10.4% depending on the microsites. The occurrence and duration of PPFD above 80 μmol m⁻² s⁻¹, which might reflect sunfleck activity, varied greatly among the microsites. However, several simple empirical relationships were found between some parameters characterizing microsite light availability and sunfleck activity; the diffuse site factor was correlated well with other parameters, including daily total PPFD, daily totals and daily summed durations of high PPFD above any examined threshold level, and its contribution to daily total PPFD. Diffuse site factors which were obtained for 700 microsites within an area of 28 m² on three different occasions during the growing season showed high correlations within the microsite. Based on the regressed relationship between the relative growth rate of current-year seedlings ofQuercus serrata and the microsite diffuse site factor and the results of area-survey measurement of the diffuse site factor, an estimation was made of the abundance of potential ‘safe-sites’ for seedling growth of the species; the ‘safe sites’ were estimated to cover 40% and 0% of the total area of the sunny and shady sites of the forest, respectively.     

Carbohydrate storage in five resprouting Florida scrub plants across a fire chronosequence

Most research analyzing nonstructural carbohydrate (NSC) concentrations on resprouter species in fire-controlled ecosystems has concentrated on how NSC concentrations recover immediately after fire. However, we know little of the effect of long periods without fire on NSC concentrations. In order to assess the effect of different periods of time-since-fire on resprouter species, we studied carbohydrate concentrations (total [NSC], soluble sugars [SS] and nonsoluble sugars [NSS]) in five resprouting species with contrasting trends of abundance across a chronosequence of time-since-fire (0.5-40 yr) in Florida. Carbohydrate concentrations were highest in species with specialized reserve organs. [SS] was mainly explained by factors related to plant size, whereas time-since-fire was the main factor explaining [NSS]. Changes in [NSS] and [NSC] were correlated with the time-since-fire abundance patterns. Variation in [NSS] carbohydrates can be related to the structural development of vegetation, with only those species capable of accessing full light able to accumulate carbohydrates, whereas subordinate plants show reductions in the [NSS] carbohydrate fractions. In areas with long intervals between fires, this carbohydrate reduction could affect subsequent postfire resprouting vigour, although this remains to be confirmed.     

Rosette scrub occurrence and fog availability in arid mountains of Mexico

Abstract. Large succulent leaf rosettes are a characteristic life form in many deserts. In certain areas they become the dominant life form, creating a vegetation type indicated as rosette scrub. The large number of rosette species suggests a close relationship between form and environment. Rosettes are excellent harvesters of low‐intensity rains and fogs. We propose that some rosette‐dominated formations of the Mexican mountains, namely the montane rosette scrub, occur in altitudinal belts around mountains where fog is abundant. We sampled four altitudinal gradients in mountains with different flora recording the abundance and richness of plants. At one site, the Tehuacán Valley, we also measured the quantity of fog along the gradient, below, above and in the rosette scrub for one year. We found that the abundance and richness of succulent rosette species are strongly associated with altitude, showing maximum values in the well‐defined elevational belts where the montane rosette scrub occurs. Other life forms, such as stem succulent cacti or woody shrubs, do not show this mid‐elevation pattern. The altitudinal ranges where the montane rosette scrub occurs usually coincide with areas where clouds and fog form. Our micrometeorological measurements indicate that rosette plants growing within a cloud belt can increase their water supply by 10–100% by harvesting fog. Outside these belts fog harvest is negligible. Desert rosettes constitute one of the most common fog‐harvesting growth forms.     

Habitat matters: The role of spore bank fungi in early seedling establishment of Florida slash pines

This study investigated broad patterns in communities of ectomycorrhizal fungi from three Florida habitats (sandhills, scrub, and pine rocklands) and the ability of spore bank fungi to associate with Pinus elliottii (slash pine) and Pinus densa (south Florida slash pine). Efforts to replant pines in the endangered pine rocklands are vital to the persistence of this habitat, yet little is known about the ectomycorrhizal fungi communities or how they may differ from those in other pine-dominated habitats in Florida. We used high-throughput amplicon sequencing (HTS) to assess baseline fungal communities and greenhouse bioassays to bait ectomycorrhizal fungi using seedlings. HTS soil data recovered 188 ectomycorrhizal species but only a few subsequently colonized the bioassay seedlings. We recovered 21 ectomycorrhizal species on pine seedlings including common spore bank fungi such as Cenococcum, Suillus, and Tuber, but Rhizopogon species were dominant across all sites and habitats. Habitat type and site were significant variables influencing the community composition of the total soil fungal community, soil ectomycorrhizal community, and the fungi found on seedling root tips. However, we found no significant differences between the ectomycorrhizal communities on seedling roots from the two Pinus species.     

Influences of neighbouring plants on seedling establishment in limestone grassland

Abstract. Influences of neighbouring plants on seedling establishment of six dicotyledonous species was investigated in a nutrient-poor limestone grassland (Mesobrometum) in northern Switzerland. Microsites with different vegetational structure were created and seeds sown in them. The fate of emerging seedlings was followed for one or two years. A plant surviving its first growing season was regarded as established. Shelter by neighbouring plants appeared to be necessary for the establishment of Arabis hirsuta and Primula veris. Because of drought and frost heave, hardly any seedlings of these species established in gaps (4 - 21 %). In microsites with vegetation, their survival was significantly higher (40–57 %). Neighbouring plants slightly reduced the survival of Plantago lanceolata and Sanguisorba minor, but these species established well both in gaps (74 - 81 %) and in vegetated microsites (54 - 67 %). Medicago lupulina established well in all microsites in one year (71 - 79 %), but poorly in the next year (18–32 %). Linum catharticum emerged poorly in one year and was completely extinguished by a fungal pathogen the following year. The overall conclusion is, that gaps are of minor importance for recruitment of these species in this grassland. Physical hazards and pathogens control seedling establishment to a greater extent than competition by neighbouring plants. Some species are hardly able to establish without shelter of vegetation. Seed size is an important factor for success of establishment, especially in gaps. It is suggested, that the relatively low productivity and the absence of litter accumulation (due to mowing and biomass removal) are important conditions for the observed behaviour of the seedlings and juvenile plants in this community.     

Elephant herbivory,frost damage and topkill in Kalahari sand woodland savanna trees

Questions: Which factors best predict the probability of elephant and frost damage in Kalahari sand woodland savanna? What is the association between tree mortality and the disturbance regime? Location: Western Zimbabwe. Methods: Elephant and frost damage, topkill, and whole‐plant mortality were quantified in ten common tree species in a Kalahari sand savanna in Zimbabwe. Individual trees were tagged in 20 plots and monitored over a two‐year period. A model selection approach was used to test the association between the probability of damage and size, prior damage, and neighbourhood effects, and to investigate the effect of damage on mortality. Results: Elephant damage differed strongly among species, and was not influenced by neighbourhood effects or prior disturbance. Frost damage also varied across species, and declined as a function of stem size and neighbourhood tree cover, and ‐ against expectations ‐ prior disturbance. Topkill increased as a function of elephant and frost damage, but was lower in previously damaged than in undamaged trees. Conclusions: Frost and elephant damage are influenced by community composition, and frost damage is also correlated with community structure and prior disturbance. Frost is an important and generally overlooked disturbance agent in southern African woodlands, where it may play a key role in association with other disturbance factors ‐ such as elephant herbivory ‐ that reduce woodland canopy cover.     

Facilitation of seedling establishment by an endemic shrub in tropical coastal sand dunes

Facilitation is predicted to occur on coastal sand dunes as these ecosystems have the harsh physical conditions common during primary succession. In a tropical dune system on the Gulf of Mexico the spatial patterns of plant distribution were analyzed and the hypothesis that facilitation is the responsible mechanism was tested by monitoring changes in the micro-environment and performing a seedling planting experiment under natural conditions. Densities of seedlings and adults of late colonizer grasses were two to six times greater beneath the early colonizer shrub, Chamaecrista chamaecristoides, than in the exposed areas. Temperatures on the sand surface, wind speed and sand accretion were significantly reduced by the shade of Chamaecrista. Only phosphate contents during the dry season were significantly higher in sand in the shade than in exposed sand. In the field experiment, successful establishment of seedlings of two late colonizer grasses (Trachypogon plumosus and Schizachyrium scoparium) was low but was concentrated exclusively beneath the shrubs. The introduced plants were reproductive one year after onset of the experiment and remained vigorous after three years. The observed spatial aggregation among the target species may be a result of improved conditions in the shade of the shrub, which facilitated the survival and establishment of late colonizers. On top of the environmental amelioration, protection from recurrent disturbances such as substrate mobility is an additional beneficial effect of the shrub.     

Seed mass versus seedling performance in Scots pine: a maternally dependent trait

It is generally accepted that larger seeds give rise to seedlings with better performance. On the other hand, the size that a seed reaches is genetically determined by at least two different traits ; the genetic variability of the developing embryo and the genetic variability of the maternal plant. Thus, the relative contributions of these two traits affect seedling performance by influencing seed size. In this paper, I investigate the effect of seed size on seedling performance in the Scots pine ( Pinus sylvestris ). From eight maternal plants, 50 seeds were planted in each of two soil types (800 seeds in total), and seedling performance was monitored for 1 yr. Seed mass proved to be highly constant within maternal plants. Soil type influenced emergence and survival; however, the effect of soil type differed depending on maternal origin. Seed mass was positively correlated with seedling emergence, although this relationship was not found for seedling survival or date of emergence. The initial growth of the shoot was also positively correlated with seed mass. However, after one growing season, seed mass had no effect on seedling performance, which depended exclusively on maternal origin. Nevertheless, the mean mass of seeds produced by plants was positively correlated with mean values of growth parameters. Thus, first-year seedling performance seems to be a maternal trait indirectly associated with seed size.     

Interspecific variation in tree seedling establishment in canopy gaps in relation to tree density

Abstract. We tested whether interspecific variation in tree seedling establishment in canopy gaps was significantly related to interspecific variation in tree density, for seven deciduous forest tree species (Quercus alba, Hamamelis virginiana, Acer rubrum, Sassafras albidum, Quercus rubra, Prunus serotina, Ostrya virginiana). For each species, seedling establishment was calculated as the difference in seedling density before experimental gap creation versus three years after gap creation. In each of the six experimentally-created gap types (33 % or 66 % removal of tree basal area from 0.01-ha, 0.05-ha or 0.20-ha patches), differences in seedling establishment among species were significantly related to differences in their density in the tree canopy. A regression model with log_e tree density as the independent variable accounted for between 93 % and 98 % of interspecific variation in seedling establishment. Our results provide empirical support for models of tree dynamics in gaps that assume seedling establishment depends on canopy tree density.     

Microhabitat associations and seedling bank dynamics in a neotropical forest

We conducted a rigorous test of tropical tree seedling microhabitat differentiation by examining microhabitat associations, survival and growth of established seedlings of ten tropical tree species representing a four-factor gradient in seed size. Eight microhabitat variables describing soil and light conditions were measured directly adjacent to each of 588 seedlings within twelve 10×100 m belt transects at Paracou, French Guiana, and at 264 reference points along the transects. From these measurements, we defined three principal components describing soil richness, soil softness and canopy openness. Six of ten species (in 9 of 30 total cases) were distributed non-randomly with respect to microhabitat along at least one principal component. However, few species demonstrated clear microhabitat specialization. All shifts in distribution relative to reference points were in the same direction (richer, softer soil). Furthermore, of 135 pairwise comparisons among the species, only 7 were significantly different. More than three-fourths of all seedlings (75.3%) survived over the 2-year monitoring period, but survival rates varied widely among species. In no case was the probability of survival influenced by any microhabitat parameter. Relative height growth rates for the seedlings over 2 years varied from –0.031 cm cm^–1 year^–1 (Dicorynia guianensis, Caesalpiniaceae) to 0.088 cm cm^–1 year^–1 (Virola michelii, Myristicaceae). In only 4 of 30 cases was height growth significantly associated with one of the three principal components. Because the conditions in this study were designed to maximize the chance of finding microhabitat differentiation among a group of species differing greatly in life history traits, the lack of microhabitat specialization it uncovered suggests that microhabitat partitioning among tropical tree species at the established seedling stage is unlikely to contribute greatly to coexistence among these species.     

Seedling establishment of deciduous trees in various topographic positions

Abstract. We investigated the effect of topography‐related environmental factors (i.e., ground‐surface stability and soil moisture) on seedling establishment of 8 deciduous tree species. A field experiment was carried out using canopy species, which were classified into 3 groups based on the spatial distribution of adult trees (ridge, slope and valley). Demographic parameters were compared among species during the early stage of seedling establishment among 3 topographic positions in combination with gap and canopy conditions. The percentage of emerging seedlings tended to be lower on the ridge irrespective of the adult topographical distribution patterns. There was no clear trend in seedling emergence among the species groups classified by their spatial distribution. Seedling survival during 2 growing seasons was significantly different among species, topographic positions and light conditions. On the ridge, seedlings of the species dominating ridge tops had greater survival than those of other species, probably due to differences in demand for soil moisture. On the slope, frequent physical damage caused by surface material movement was observed and some species showed greater adaptability to the disturbed slope habitat. Survival of all seedlings was highest in the valley plots. Light conditions were the critical factor for seedling survival in some species. The results of this study suggest that topography creates diverse habitats for the establishment of tree seedlings. In addition to soil moisture, surface material movement may be a significant factor affecting seedling establishment.     

Conifer seedling recruitment in a southeastern Canadian boreal forest: the importance of substrate

Abstract. In order to explain conifer species recruitment in Canada's southeastern boreal forest, we characterized conifer regeneration microsites and determined how these microsites vary in abundance during succession. Microsite abundance was evaluated in deciduous, mixed and coniferous stands along a 234-yr postfire chronosequence. Conifers were most often found in relatively well-illuminated microsites, devoid of litter, especially broad-leaf litter, and with a reduced cover of lower vegetation (< 50 cm tall). Although associated with moss-rich forest floor substrates, Abies balsamea was the most ubiquitously distributed species. Picea glauca and especially Thuja occidentalis seedlings were frequently found on rotten logs. Light measurements did not show differences among seedling species nor between stand types. The percentage cover of broad-leaf litter decreased significantly during succession. Also, rotten logs covered with moss occupied a significantly larger area in the mid-successionnal stands than in early successional deciduous or late successional coniferous stands. The results suggest that the presence of specific forest floor substrate types is a factor explaining low conifer recruitment under deciduous stands, conifer codominance in the mid-successional stage, and delayed Thuja recolonization after fire. Results also suggest that some facilitation mechanism is responsible for the observed directional succession.     

The spatial characteristics of stand structure in Pinus torreyana

The arrangement of trees within a stand by location and age (stand structure) is in part determined by the life history strategy of the species and the disturbance history of the stand. In western North America such disturbances are often the product of wildfires and human management activities. The current study uses spatial analysis to characterize three stands of Pinus torreyana with known disturbance histories. Two stands are located at Torrey Pines State Reserve (TPSR). Of these, one stand has burned twice since 1972. Fire has been successfully excluded from the other stand since the early part of this century. A third stand, on Santa Rosa Island, Channel Islands National Park (SRI), has been grazed heavily since the mid-19th century and has not experienced fire since that time. One-hectare study plots were established in the interior of each stand. Considering the known life history attributes of P. torreyana, and the disturbance histories of the stands, predictions are made concerning the spatial characteristics of the respective stands. All trees within each study plot were sized by diameter breast height (DBH) and mapped. Three techniques of spatial analysis are applied to the resulting unweighted point pattern distributions and the distributions weighted by the square root of DBH as a surrogate for age. The results are consistent with predictions and confirm the following generalizations concerning patterns of aggregation in Torrey pine stands. Young trees tend to be more aggregated than old trees within the same stands. Young stands tend to be more aggregated than old stands on otherwise ecologically similar sites. On a periodically disturbed site there are clusters of trees that represent cohorts of post-disturbance recruitment.     

The role of in vitro cultivation on symbiotic trait and function variation in a single species of arbuscular mycorrhizal fungus

In vitro propagation of AM fungi using transformed root cultures (TRC) is commonly used to obtain pure AM fungal propagules for use in research and industry. Early observations indicate that such an artificial environment can alter traits and function of AM fungi over time. We hypothesized that increased in vitro cultivation may promote ruderal strategies in fungi by enhancing propagule production and reducing mutualistic quality. To examine the effect of in vitro cultivation on the trait and function of AM fungi, we inoculated plants with 11 Rhizoglomus irregulare isolates which fell along a cultivation gradient spanning 80 generations. We harvested plants at 10, 20 and 30 d post inoculation to observe differences in fungal and plant traits post infection. In vitro cultivation led to increased spore production but reduced plant shoot phosphorus. Our results indicate that in vitro propagation may indirectly select for traits that affect symbiotic quality.