Responses of plant and bird communities to prescribed burning in tallgrass prairies

Historic losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to declines in diversity and abundance of plants and birds associated with such habitat. Prescribed burning is a management strategy that has potential for restoring and rejuvenating prairies in fragmented landscapes, and through such restoration, might create habitat for birds dependent upon prairies. To provide improved data for management decision-making regarding the use of prescribed fire in tallgrass prairies, we compared responses of plant and bird communities on five burned and five unburned tallgrass prairie fragments at the DeSoto National Wildlife Refuge, Iowa, USA, from 1995 to 1997. Overall species richness and diversity were unaffected by burning, but individual species of plants and birds were affected by year-treatment interactions, including northern bobwhite (Colinus virginianus) and ring-necked pheasant (Phasianus colchicus), which showed time-delayed increases in density on burned sites. Analyses of species/area relationships indicated that, collectively, many small sites did make significant contributions to plant biodiversity at landscape levels, supporting the overall conservation value of prairie fragments. In contrast, most birds species were present on larger sites. Thus, higher biodiversity in bird communities which contain area-sensitive species might require larger sites able to support larger, more stable populations, greater habitat heterogeneity, and greater opportunity for niche separation.     

Fire,canopy cover and seedling dynamics in Mediterranean shrubland of northeastern Spain

Abstract. Seedling establishment and survival of the main woody species of early-successional shrubland in northeastern Spain were studied from 1992 to 1995 with emphasis on the importance of vegetation cover (existence of open areas and the situation beneath the plant canopy) and microhabitat (occurrence of stones, litter and bare soil). In the absence of fire, vegetation cover (measured at a scale of 30cm × 30 cm) was not correlated with seedling emergence of most species, nor with growth and survival of seedlings up to one year old. Seedlings older than one year showed a similar pattern: their density was not significantly different in both kinds of habitats. The emergence of seedlings was mainly associated with the presence of mature plants – which can provide seeds – and with the absence of unsuitable microhabitats (large stones, deep litter and bare soil). In April 1994 a wildfire burned the study area. This enabled a study of the pattern of post-fire establishment. After the wildfire, seedling emergence of several species increased and most species produced seedlings with higher survival and growth rates than in the period before the fire. Pre-fire cover, however, did not show significant effects on post-fire seedling dynamics in most cases.     

The role of nurse functional types in seedling recruitment dynamics of alternative states in rangelands

In arid ecosystems, recruitment dynamics are limited by harsh environmental conditions and greatly depend on the net outcome of the balance between facilitation and competition. This outcome can change as a consequence of degradation caused by livestock overgrazing. Also, distinct plant species may show a differential response to a common neighbour under the same environmental conditions. Therefore, ecosystem degradation could affect the net balance of plant-plant interactions, which can also depend on the functional traits of potential nurse species. The aim of this study is to assess the influence of alternative degradation states on (i) the density of seedlings of perennial species emerging in four microsite types, and on (ii) the relative interaction intensity (RII) between seedlings and potential nurses belonging to three functional types (deep- and shallow-rooted shrubs, and tussock grasses). During three years, we recorded seedling density of perennial species in four alternative degradation states in grass-shrubby steppes from northwestern Patagonia. The density of emerged seedlings of perennial species decreased sharply as degradation increased, showing non-linear responses in most microsites. Seedling density underneath deep-rooted shrubs was higher than underneath shallow-rooted shrubs and tussock grasses. Also, deep-rooted shrubs were the only functional type that recorded seedling emergence in highly degraded states. Deep-rooted shrubs had facilitative effects on the seedlings emerging and surviving underneath them, independently of ecosystem degradation. In contrast, RII between shallow-rooted shrubs and recently emerged seedlings, switched from positive effects in the less degraded states, to negative effects in the most degraded state. Tussock grasses recorded the weakest intensity of facilitative interactions with recently emerged seedlings, switching to competitive interactions as degradation increased. Our results suggest that species with key functional traits should be considered in management and restoration plans for rangelands with different degradation levels, since they have a strong influence in the net outcome of plant-plant interactions and in the recruitment dynamics of arid ecosystems.     

Shrub emergence and seedling growth in microenvironments created by Prosopis glandulosa

Abstract. Facilitation of the establishment of certain plant species by nurse plants is a common phenomenon in arid and semiarid ecosystems. The most commonly reported mechanisms of facilitation include cooler temperatures and increased soil nutrients beneath the nurse plant canopy, which favor establishment of other plant species. During conversion of upland grasslands to thorn woodland in southern Texas, Prosopis glandulosa appears to facilitate establishment of other woody plants, including Celtis pallida, whereas Acacia smallii occurs only in habitats between P. glandulosa canopies. We tested the hypothesis that light intensity and soils under P. glandulosa canopies facilitate seedling emergence and growth of C. pallida but inhibit seedling emergence and growth of A. smallii. In the field, C. pallida and A. smallii seeds were planted under P. glandulosa canopies and in adjacent interspaces. Percent emergence of C. pallida seedlings was greater under the canopy of P. glandulosa, whereas percent emergence of A. smallii seedlings was greater in interspaces. In a greenhouse experiment, seeds of each species were planted in pots filled with soil from under P. glandulosa canopies or from adjacent interspaces. Two treatments, shade and sunlight, were imposed and plants harvested seven weeks later. Seedling mass of both species was greater in canopy soil than in interspace soil in sunlight but mass of the two species did not differ between soil sources in shade. Canopy soils contained more total and available nitrogen than interspace soils. These results suggest that light is more limiting than nutrients under shaded conditions and so neither species can take advantage of the high nutrients beneath P. glandulosa. Shade and greater soil nutrients beneath P. glandulosa do not appear to be the major factors that facilitate C. pallida or inhibit A. smallii. Aggregation of C. pallida beneath P. glandulosa canopies appears to be a complex process that involves both passive facilitation (seed dispersal by birds) and active facilitation (reduction of seed dormancy by under-canopy temperatures) operating only during the seed germination stage with successional mechanisms other than facilitation operating during later stages of shrub establishment and growth.     

Rodent seed predation and seedling recruitment in mesic grassland

Seedling recruitment of two grasses (Arrhenatherum elatius and Festuca rubra) and two herbs (Centaurea nigra and Rumex acetosa) was measured in areas with and without rodents to which seeds of each species were sown at three seed densities (1000, 10,000 and 50,000 seeds m⁻²) in two seasons (spring and autumn 1995). Seed removal was measured for 10-day periods and the fate of seedlings was followed for 15 months after sowing. The proportion of seed removed ranged from 6 to 85% and increased with increasing seed density for each species. Rodents had no effect on seedling emergence or survival in the spring sowing. In the autumn sowing, rodents reduced seedling emergence of all four species sown at 1000 and 10,000 seeds m⁻² but had no impact at 50,000 seeds m⁻², presumably because of microsite limitation. We suggest the difference between spring and autumn arose because emergence was seed limited in autumn but microsite limited in spring; microsite availability was higher in autumn because a summer drought killed plants, reduced plant biomass and opened up the sward. Fifteen months after the autumn sowing, fewer A. elatius and C. nigra seedlings survived on plots exposed to rodents. This result reflected not only the reduced seedling emergence but also increased seedling mortality (seedling herbivory) in sites exposed to rodents. In contrast, F. rubra and R.acteosa showed density-dependent seedling survival which compensated for initial differences in seedling emergence, so that no effect of rodents remained after 15 months. The results suggest that rodent seed predation and seedling herbivory exert strong effects on seedling recruitment of A.elatius and C. nigra when recruitment conditions are favourable (conditions that lead to high microsite availability) and may contribute to both species being maintained at low densities in the grassland. The results also demonstrate that highly significant impacts of rodent seed predation at the seedling emergence stage can disappear by the time of plant maturation. Received: 2 March 1998 / Accepted: 28 September 1998     

Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands

The season in which a fire occurs may regulate plant seedling recruitment because of: (1) the interaction of season and intensity of fire and the temperature requirements for seed release, germination and growth; (2) post-fire rainfall and temperature patterns affecting germination; (3) the interaction of post-fire germination conditions and competition from surrounding vegetation; and (4) the interaction of post-fire germination conditions and seed predators and/or seedling herbivores. This study examined the effects of different fire intensities and fire seasons on the emergence and survival of shrubs representing a range of fire response syndromes from a summer rainfall cool climate region. Replicated experimental burns were conducted in two seasons (spring and autumn) in 2 consecutive years and fuel loads were increased to examine the effects of fire intensity (low intensity and moderate intensity). Post-fire watering treatments partitioned the effects of seasonal temperature from soil moisture. Higher intensity fires resulted in enhanced seedling emergence for hard-seeded species but rarely influenced survival. Spring fires enhanced seedling emergence across all functional groups. Reduced autumn recruitment was related to seasonal temperature inhibiting germination rather than a lack of soil moisture or competition. In Mediterranean-type climate regions, seedling emergence has been related to post-fire rainfall and exposure of seeds to seed predators. We think a similar model may operate in temperate summer rainfall regions where cold-induced dormancy over winter exposes seeds to predators for a longer time and subsequently results in recruitment failure. Our results support the theory that the effect of fire season is more predictable where there are strong seasonal patterns in climate. In this study seasonal temperature rather than rainfall appears to be more influential.     

Genecology of Douglas fir in western Oregon and Washington

BACKGROUND AND AIMS: Genecological knowledge is important for understanding evolutionary processes and for managing genetic resources. Previous studies of coastal Douglas fir (Pseudotsuga menziesii var. menziesii) have been inconclusive with respect to geographical patterns of variation, due in part to limited sample intensity and geographical and climatic representation. This study describes and maps patterns of genetic variation in adaptive traits in coastal Douglas fir in western Oregon and Washington, USA. METHODS: Traits of growth, phenology and partitioning were measured in seedlings of 1338 parents from 1048 locations grown in common gardens. Relations between traits and environments of seed sources were explored using regressions and canonical correlation analysis. Maps of genetic variation as related to the environment were developed using a geographical information system (GIS). KEY RESULTS: Populations differed considerably for adaptive traits, in particular for bud phenology and emergence. Variation in bud-set, emergence and growth was strongly related to elevation and cool-season temperatures. Variation in bud-burst and partitioning to stem diameter versus height was related to latitude and summer drought. Seedlings from the east side of the Washington Cascades were considerably smaller, set bud later and burst bud earlier than populations from the west side. CONCLUSIONS: Winter temperatures and frost dates are of overriding importance to the adaptation of Douglas fir to Pacific Northwest environments. Summer drought is of less importance. Maps generated using canonical correlation analysis and GIS allow easy visualization of a complex array of traits as related to a complex array of environments. The composite traits derived from canonical correlation analysis show two different patterns of variation associated with different gradients of cool-season temperatures and summer drought. The difference in growth and phenology between the westside and eastside Washington Cascades is hypothesized to be a consequence of the presence of interior variety (P. menziessii var. glauca) on the eastside.     

Fire and Litter Effects on Seedling Establishment in Western Oregon Upland Prairies

Prescribed burning is an important tool for managing and restoring prairies and other ecosystems. One effect of fire is plant litter removal, which can influence seedling establishment. Four experimental treatments (burned, clipped and raked to remove litter, burned with litter reapplied, and unmanipulated) were applied to 2 × 2.5–m plots in three western Oregon, United States, upland prairies to determine how burning affects seedling establishment. Seeds of common exotic and native prairie species were sowed into the experimental plots after treatments. Seedlings were censused the following spring. The experiment was repeated on each of the three sites, representing three common types of prairie vegetation: an Annual Exotic Grass site, a Perennial Exotic Grass site, and a Native Bunchgrass site. In both the Annual Exotic Grass and the Perennial Exotic Grass sites, burning significantly improved native, but not exotic, seedling establishment over those on unburned plots. Litter removal was a significant component of this burn effect, particularly on the Perennial Exotic Grass site. In these winter‐moist systems, the net effect of litter is to inhibit seedling establishment. Burning treatments on the Native Bunchgrass site significantly increased seedling establishment only of short‐lived exotic species. These results suggest that in prairie ecosystems similar to the Annual and Perennial Exotic Grass sites, prescribed burning followed by sowing native seeds can be an effective restoration technique. Burning alone or sowing alone would be counter‐productive, in the first case because increased establishment would come from exotic species and in the second case because establishment rates are low in unburned plots.     

沙埋对六种沙生植物种子萌发和幼苗出土的影响

研究了沙埋对科尔沁沙地6种优势植物的种子萌发和幼苗出土的影响．进行0．2、4、6、8、10和12cm等7个深度沙埋处理．结果表明,在不同沙埋处理时,沙蓬萌发差异显著。而差不嘎蒿2锄埋深与其他埋深的发芽差异显著,其他4种植物0cm埋深与其他埋深的发芽差异显著;沙埋对所有植物幼苗出土均有显著影响,埋深增加,出苗率减小;繁殖体大的物种与繁殖体小的物种相比,能从更深沙层中出苗,幼苗出土最大深度排序为苦参>东北木蓼≥沙蓬>山竹子>雾冰藜>差不嘎蒿．     

Effect of grass litter on seedling recruitment of the critically endangered Cistus heterophyllus in Spain

Cistus heterophyllus Desf. is an Iberian-north African endemic plant species with only one wild population left in Spain. A sowing experiment under controlled conditions was designed to assess both the seedling emergence rate and the growth fitness of C. heterophyllus on a litter layer of Brachypodium retusum (Pers.) Beau., which forms dense carpets around the remnant population of C. heterophyllus. In order to test the physical effects of B. retusum litter on Cistus seedling recruitment, six sowing treatments in pots were compared that combined three doses of litter (0, 0.6 and 1.1 kg m⁻²) and different physical locations of the seeds (above, below the litter layer or within a litter–peat mixture). B. retusum litter interfered with the seedling recruitment of C. heterophyllus, negatively affecting both seedling emergence and their early growth. Such an effect was more severe when the seeds were located above a 0.6 kg m⁻² litter layer, although seedlings from seeds sown under the litter showed a growth reduction ranging from 58% to 100% compared to the no-litter treatment. A significant negative effect of the litter–peat mixture on the number of leaves, leaf size and number of branches of seedlings was also inferred. These data provide experimental evidence on the serious threat that B. retusum litter represents for the conservation and regeneration of the last population of C. heterophyllus in Spain.     

Fine root production and carbohydrate concentrations of mature longleaf pine (Pinus palustris P. Mill.) as affected by season of prescribed fire and drought

The historical range of longleaf pine (Pinus palustris P. Mill) has been greatly reduced, in part, by lack of fire. Recently, the application of fire has become an accepted practice for the restoration of longleaf pine to former parts of its natural range. This study was designed to evaluate the effects of season of prescribed fire on the root growth and nonstructural carbohydrate concentrations of longleaf pine, and identify the time of year when fire has the least negative effect on longleaf pine root processes. We found that root growth was generally less on July-burned plots than on either March- or May-burned plots and we attribute these responses to the effect of fire on interaction between root processes and the soil environment. Specifically, soil water and temperature conditions may have been less favorable for root growth on the July-burned plots compared to the March- and May-burned plots. With two years of information on the seasonal dynamics of foliage production, root growth, and root carbohydrates, we determined that at our study site, optimal prescribed fire would impact tree growth less in November through March compared to other times of the year. We also observed that severe drought during the 1998 growing season was associated with a 3-month delay in peak root growth, and prolonged drought in 1999 through 2000 coincided with a reduction in root starch storage. We conclude that season of prescribed fire potentially affects root processes, but that severe or prolonged drought may either interact with or override these effects.     

Effects of sand burial depth and seed mass on seedling emergence and growth of Nitraria sphaerocarpa

A greenhouse experiment was conducted to test the effects of sand burial depth and seed mass on seedling emergence and growth of Nitraria sphaerocarpa. Seeds of Nitraria sphaerocarpa were sorted into three size-classes (large, medium, small) and artificially buried at 0, 1, 2, 3, 4, 5 and 6 cm depths in plastic pots filled with unsterilized sand. In the seven treatments, the percent emergence, seedling mass and seedling height, significantly affected by both burial depth and seed size, were highest at the optimal burial depth of 2 cm burial depth, and decreased with increasing burial depth in each seed size-class. Although seedling mass was usually greatest for large seeds and least for small seeds at each burial depth, little difference was observed in seedling height at shallow burial depths of 0–3 cm. In each seed size-class, with increasing burial depth, both root-mass ratio and aboveground stem-mass ratio decreased, while belowground stem-mass ratio increased. In each burial depth, with decreasing seed size, belowground stem-mass ratio increased, while root-mass ratio decreased.     

Quercus rugosa seedling dynamics in relation to its re-introduction in a disturbed Mexican landscape

Abstract. In order to define the ‘regeneration niche’ and then promote the re-introduction of Quercus rugosa into a disturbed area we studied seed predation, germination, seedling survival and growth of this species at three different sites: the forest interior, the forest border and a disturbed site, along a disturbance gradient. Acorn removal on the ground was high at the sites, with higher removal rates from 25-seed clusters than from 5- and 1-seed clusters. More seeds were removed at the forest border than at the other two sites, although in all cases some seeds remained in the soil for periods long enough for them to germinate and produce seedlings. Seed germination and seedling establishment were more successful and similar in the forest interior and the forest border, and much less successful at the disturbed site, due to acorn desiccation. Seedling survival was strongly affected by site, with the highest survival at the forest border and the lowest survival at the disturbed site after a year. The subsequent dry season, which was particularly harsh, caused a drop in survival both at the forest border and the disturbed site. In the former, a closer analysis revealed that partially shaded microsites enhanced survival, while in the latter overall survival was very low, with approximately 5 % of the seedlings surviving, compared to 50 % final survival at the other sites. There was a positive correlation between seed size and seedling survival at the forest border. The three size variables measured (seedling height, basal diameter and crown area) did not increase noticeably during the 20-month study period at the forest interior, while growth still occurred at the forest border and the disturbed site. However, during the two dry seasons there were large drops in mean seedling height and crown area at the disturbed site, due to more frequent dieback, causing large variation in seedling size. These results show that extensive re-introduction of this species to the disturbed area will probably fail under present conditions, and that careful selection or duplication of microsites that mimic the partial shade of the forest border is a sound method for improving oak establishment at this site.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Fire season affects size and architecture of Colophospermum mopane in southern African savannas

Wildfires may be started naturally by lightning or artificially by humans. In the savanna regions of southern Africa, lightning fires tend to occur at the start of the wet season, during October and November, while anthropogenic fires are usually started during the dry season, between July and August. A long-term field manipulation experiment initiated in the Kruger National Park in 1952 was used to explore whether this seasonal divergence affects tree abundance, spatial pattern, size and architecture. After 44 years of prescribed burning treatments that simulated the seasonal incidence of lightning and anthropogenic fires, mean densities of the locally-dominant shrub, Colophospermum mopane, were 638 and 500 trees ha^–1 respectively. Trees in burnt plots had aggregated distributions while trees in unburnt plots had random distributions. Significant differences (p < 0.001) were recorded in a range of morphological parameters including tree height, canopy diameter, mean stem circumference and number of stems. The incidence of resprouting also differed significantly between treatments, with burnt trees containing a high proportion of coppiced stems. The differences in tree size and architecture between the mid-dry season and early-wet season burning plots suggest that anthropogenic fires applied during July and August cannot substitute for a natural lightning fire regime. Anthropogenic fire yields a landscape that is shorter, more scrubby and populated by numerous coppiced shrubs than the landscape generated by natural lightning fire conditions.     

Optimal time of seedling emergence in a dune-population ofOenothera glazioviana

Oenothera glazioviana (= O. Erythrosepala) behaves as a facultative biennial in an infertile sand-dune system in Japan. In the field, seedlings emerge mainly in autumn following seed ripening in late summer with an additional small peak in spring. The effects of emergence time on subsequent survival and fecundity, and the ultimate numbers of seeds produced per emerging seedling (F _s) were examined by a demographic model. The model showed that seedlings emerging either in March or in September–October will attain higherF _s than those emerging in other months. If mortality in the seed bank is taken into account, autumn seems to be the best season for emergence to maximize the number of seedlings in the next generation.     

Spatial and environmental factors contributing to patterns in arboreal and terrestrial oribatid mite diversity across spatial scales

Recruitment is a crucial event in the plant life cycle that is very sensitive to interaction with established vegetation. Based on a large comparative experiment, we tested the hypothesis that the components of recruitment––emergence time and rate, seedling survival and biomass––differ in response to plant–plant interactions during recruitment. The consequences for the population are predicted with a simple demographic model assessing the response of seed production. In a common garden experiment, we recorded the recruitment of four target species in an individual-based survey protocol. A total of 7,680 seeds were sown within 20 neighbourhoods, consisting of 19 mono-specific herbaceous stands and a control treatment without vegetation. We measured transmitted light, temperature and moisture at soil surface to characterise the environmental conditions within neighbourhoods. The mean height of neighbours controlled temperature buffering and light interception and thus depicted the interaction gradient. Emergence rate and time increased with neighbour height in two of the four target species, while seedling survival and biomass significantly decreased with neighbour height in three and all four target species, respectively. We recorded a shift in seedling neighbour interactions under the tallest neighbours that largely favoured emergence but strongly depressed seedling survival and biomass. The components of recruitment were predicted to differ in their impact on later adult performance. Biomass strongly contributed to predicted seed production in three target species, and emergence had an equal or greater impact on a fourth species. These results confirm the fundamental role of plant–plant interactions in the recruitment of herbaceous species through a complex combination of habitat amelioration, which facilitates emergence and light competition, which in turn limits seedling survival and biomass.