Secondary succession after perturbations in a shrubland community

Shrubland responses to experimental burning, cutting and ploughing treatments were studied over 15 years in two shrubland communities dominated by Erica australis. The treatments represent the most frequent forms of anthropogenic disturbances experienced by these communities throughout their history. The response to burning and cutting treatments is similar, and the succession process is characterised as autosuccession. The highest values for herbaceous annuals and perennials were observed in the third and fourth years. Generally, herbaceous species remain present throughout the study period, while woody taxa ones increase their cover values over time. The quantity of herbaceous species present is in inverse proportion to the quantity of woody taxa. The woody species that appear immediately after treatments are sprouting species, namely Erica australis and Arctostaphylos uva-ursi. The response to ploughing is slower, reflecting the recovery mechanism (seedlings). However, after 15 years, there are no significant differences in regeneration between treatments. The first stages of this post-ploughing succession are dominated by annual species until the fourth or fifth years, after which woody species begin to dominate and herbaceous taxa decrease considerably. Woody species with high germination values are Halimium alyssoides and Halimium umbellatum. These shrubland communities have a very high resilience to such perturbations and start regenerating rapidly, reaching the original state in about 9 years. The appearance of the climax arboreal species of the area, Quercus pyrenaica, when it comes from germination, occurs 15 years after the perturbations.     

Recovery after Experimental Cutting and Burning in Three Shrub Communities with Different Dominant Species

The aim of this study is to compare the recovery dynamics in three shrub communities subjected to experimental burning and cutting, and situated on an altitudinal gradient. Climatic features are different in each area, but all had the common characteristic of very homogeneous vegetation cover before the disturbances, with only one shrub species clearly dominant, a different taxon in each area, and with different regeneration strategies. The first area was a heathland dominated by Calluna vulgaris, situated at an altitude of 1600 m, with a continental climate (mean annual precipitation 1320 mm). The second area was a heathland dominated by Erica australis, located at an altitude of 1000 m (mean annual precipitation 840 mm). The third area was a Cistus ladanifer shrubland, located at 900 m altitude, with a Mediterranean climate similar to that of the previous area, but with lower mean annual precipitation (470 mm). Erica australis recovers by vegetative resprouting, but Cistus ladanifer is an obligate seeder, as is Calluna vulgaris in these areas. Each experimental disturbance was carried out over 100 m² in each area. Post-fire recovery is faster in Cistus ladanifer: 2 years after burning there was 40% cover vs. less than 20% in the other two species. However, recovery after cutting was similar for Cistus ladanifer and Erica australis. Calluna vulgaris recovers very slowly, with cover values below 20% even 10 years after both disturbances. Cover of dominant shrub species is negatively correlated with cover of herbaceous species. So different recovery of dominant species lead a different community dynamic in each area.     

Interactions between mediterranean shrub species eight years after experimental fire

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

Responses of heather moorland and Mediterranean mouflon foraging to prescribed-burning and cutting

We assessed the effects of prescribed burning and cutting on mouflon (Ovis gmelini musimon × Ovis sp.) spring habitat using an experimental design (17.28 ha) of 2 burned, 2 cut, and 2 untreated plots within a homogeneous stand dominated by heather (Erica cinerea and Calluna vulgaris). Overall, we found a shift in treated plots from ligneous species to herbaceous species with high digestive and energetic values for mouflon. We also found a consistently higher number of mouflon feeding on these treated habitats compared to untreated plots. Such effects were still apparent 4 years after habitat modifications. Our approaches could be used by managers to improve and maintain the range of mouflon populations experiencing habitat loss (e.g., woody plant encroachment) and for which the condition of an animal has often a high economical value through trophy hunting. © 2011 The Wildlife Society.     

Seed banks of Erica australis and Calluna vulgaris in a heathland subjected to experimental fire

Abstract. Soil samples were collected before and after an experimental fire on a heathland in the province of León (Spain). The seed banks were assessed by counting the numbers of seedlings appearing of Erica australis and Calluna vulgaris. A total of 2285 germinable seeds/m² before burning and 1177 germinable seeds/m² after the fire were estimated for Erica australis, and 90 and 690 germinable seeds/m², respectively, for Calluna vulgaris. After the fire the number of germinable seeds/m² of Erica australis had decreased, whilst there was a considerable increase in the number of germinable seeds/m² of Calluna vulgaris. Neither seedlings nor sprouts of Calluna vulgaris had appeared in the field plots 10 months after the fire. Erica australis did recover mainly by sprouting after fire.     

Winter Burning and the Reduction of Cornus sericea in Sedge Meadows in Southern Wisconsin

To control shrubs, which are increasing in dominance in wetlands worldwide, winter burning may be an important tool, especially from the perspective of minimizing urban hazards. The goal of this project was to determine if winter burning was successful in reducing the dominance (mean percentage cover and maximum height) of Cornus sericea in sedge meadows in southern Wisconsin, where shrubs proliferated after cattle were excluded. Experimental burn and control plots were set up within sedge meadows, including an ungrazed “reference” site that had been little, if ever, grazed and a “historically grazed” site, a recovery site that had not been grazed by cattle since 1973. None of the dominant species including C. sericea was significantly affected by burning for either mean percentage cover or maximum height (analysis of variance: no burning × species interaction). Both mean percentage cover and maximum height were only weakly related to burning (28.1 and 14.3% of the variability contributed to the cumulative percentage of the coefficient of determination, respectively) at both sites based on non‐metric multidimensional scaling analysis. Although species richness increased in burned plots in 1999 and 2000, no differences were apparent between pre‐burned and unburned plots in 1997 and unburned plots in 1999 and 2000 (analysis of variance: year × burning interaction). After burning in the ungrazed site, herbaceous species appeared that had not been detected for decades, including Chelone glabra and Lathyrus palustris. Exotic species were present in both the ungrazed reference and recovery site. Although winter burning treatments did not reduce the dominance of woody shrub species in the site recovering from cattle grazing, burning was useful in stimulating the maintenance of species richness in the ungrazed sedge meadow.     

Plant–community responses to shrub cover in a páramo grassland released from grazing and burning

Shrub encroachment can follow grazing or burning release in páramo grasslands. While encroachment decreases herbaceous species richness in some grassland systems, the effects of this process on the herbaceous community in páramo grasslands are currently unknown. We collected data on shrub cover, herbaceous‐species cover and species composition in a páramo grassland 12 years after release from burning and cattle grazing near Zuleta, Ecuador. Topographic and soil measures were also included as predictor variables of differences in community composition. Contrary to studies in other systems, shrub cover did not have a significant effect on herbaceous‐species richness, whereas shrub‐species richness significantly increased with shrub cover. However, shrub cover was associated with significant shifts in herbaceous–community composition. Most notably, there was an increase in some shade‐tolerant forbs and tall‐statured wetland grasses with increasing shrub cover, and a corresponding decrease in some short‐statured grasses and early successional forbs. These results could indicate that the ameliorative effects of shrubs (e.g. frost and wind protection) in harsh alpine environments may partially compensate for the expected competitive effect of shrubs due to shading.     

Short-term study of effects of fertilisation and cutting treatments on the vegetation dynamics of mountain heathlands in Spain

The influence of management and nutrient availability on the vegetation dynamics of heathlands characterised by Calluna vulgaris and Erica tetralix were studied in three mountain sites in Northern Spain. A total of 90 plots (1 m² each) received different combinations of cutting and twice the estimated background atmospheric deposition of nitrogen (56 kg ha⁻¹ yr⁻¹). One of the two dominant ericaceous species was selectively cut by hand at ground level and their regeneration compared in the presence or absence of the other. The results after 2 years showed significant effects of the fertiliser on the vegetation cover, mainly by favouring perennial herbaceous species. There were less noteworthy effects on the number of flowers and on the annual growth of the ericaceous species. It is concluded that, in the short term, increased nutrients alone, at twice the estimated current atmospheric deposition for the area, will not alter significantly the composition of the mountain heathlands. However, once the stands reach the mature phase, the capacity of the community to regenerate after a severe disturbance diminishes. A drastic impact, such as cutting may not result in re-growth of the same shrub species but in replacement by herbaceous species, which will also benefit from the increased nutrients.     

Response of plant species and life form diversity to variable fire histories and biomass in the jarrah forest of south‐west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light. We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south‐west Australia. Fire frequency was defined as the total number of fires over a 30‐year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency.     

Plant diversity patterns in the Aljibe Mountains (S. Spain): a comprehensive account

The Aljibe Mountains are located in the southern tip of the Iberian Peninsula and have a remarkable biogeographical interest. The complete plant species list (trees, climbers, shrubs, perennial and annual herbs, ferns, lichens, bryophytes and macroscopic algae) was recorded in four 0.1 ha plots from each of the most representative community types (Quercus suber woodland, Q. canariensis forest, open heathland and Q. coccifera shrubland). Up to 119 plant species were found in total in the Q. suber woodland plot. The diversity of woody plants was analysed from 44 samples of cover (100 m line), and the herbaceous layer was explored in 200 quadrats (of 0.5 × 0.5 m). Three biodiversity components (species richness, endemism, and taxonomic singularity) were evaluated in both shrub and herbaceous layers. Open heathlands showed the highest richness of endemic species, both woody and herbaceous. The highest number of woody species was found in the evergreen Q. suber woodland, and of herbaceous species in the semi-deciduous Q. canariensis woodland. Taxonomic singularity was higher in Q. canariensis woodlands and Q. coccifera shrublands for woody species, but there were no significant differences in the herbaceous layer. Local species diversity of heathlands in this region resembles that of South African heathlands (fynbos), despite the obvious geographic and floristic distance, and contrasts with the low diversity of biogeographically closer, European temperate heathlands. The Aljibe Mountains show high diversity values for different life forms (from trees to mosses) and spatial scales (from community to region), and are rich in endemic species. Thus, this area should be recognised as a relevant unit within the Mediterranean plant diversity hot spots.     

Eleven years of recovery dynamic after experimental burning and cutting in two Cistus communities

Recovery after cutting and burning in two series of experimental shrubland plots dominated by Cistus laurifolius and Cistus ladanifer, respectively, was studied over 11 years. The two communities tend to recover by an autosuccession process after disturbances, with Cistus species being dominant from the first or second year. The greatest diversity is recorded in the first few years after the disturbances, with higher species richness, basically herbaceous ones, and lower dominance effect. Cover by woody species tends to increase but from the fourth year the differences are not significant. The highest herbaceous cover is after one year and the peak is more marked in the cut plots. Herbaceous species cover decreases afterwards and there are small changes in the different years but without clear trends. When all the samplings are compared by a TWINSPAN, the first division separates both communities and the  second division the initial and the final phases from each community. The separation between cut and burnt plots is only observed in the final phases.     

Effects of Prescribed Fire on Shinnery Oak ( Quercus havardii) Plant Communities in Western Oklahoma

Changes in structural and compositional attributes of shinnery oak (Quercus havardii Rydb.) plant communities have occurred in the twentieth century. These changes may in part relate to altered fire regimes. Our objective was to document effects of prescribed fire in fall (October), winter (February), and spring (April) on plant composition. Three study sites were located in western Oklahoma; each contained 12, 60 × 30‐m plots that were designated, within site, to be seasonally burned, annually burned, or left unburned. Growing season canopy cover for herbaceous and woody species was estimated in 1997–1998 (post‐treatment). At one year post‐fire, burning in any season reduced shrub cover, and spring burns reduced cover most. Winter and annual fires increased cover of rhizomatous tallgrasses, whereas burning in any season decreased little bluestem cover. Perennial forbs increased with fall and winter fire. Shrub stem density increased with fire in any season. Communities returned rapidly to pre‐burn composition with increasing time since fire. Fire effects on herbaceous vegetation appear to be manifested through increases in bare ground and reduction of overstory shrub dominance. Prescribed fire can be used as a tool in restoration efforts to increase or maintain within and between community plant diversity. Our data suggest that some plant species may require or benefit from fire in specific seasons. Additional research is needed to determine the long‐term effects of repeated fire over time.     

The effects of thermal scarification and seed storage on germination of four heathland species

The germination responses of 4 shrubs, Erica arborea, E. australis and Daboecia cantabrica (Ericaceae) and Genistella tridentata (Papilloneaceae) in relation to variable light and temperature regimes was determined for freshly harvested and 1 year-old seeds. Temperature treatments consisted of 5 levels of heat from 60 °C to 140 °C and two levels of exposure time, 2 and 4 min. We also examined the effects of stratification on germination of Erica arborea and E. australis. The seeds of Erica arborea were the first to begin germinating and the freshly collected ones reached the highest level of germination with light. The seed viability of E. arborea was affected by storage, decreasing germinative capacity, while mean germination in Erica australis ssp. aragonensis was lowest. However, the stored E. australis seeds germinated better than the fresh ones. Germination response to ageing varies in Daboecia cantabrica depending on heat pretreatment. A significantly positive effect of high temperature and exposure time on germination was shown in Daboecia cantabrica and Genistella tridentata. Cold pre-treatment of seeds did not enhance germination.     

Using consecutive prescribed fires to reduce shrub encroachment in grassland by increasing shrub mortality

Woody plant encroachment into open grasslands occurs worldwide and causes multiple ecological and management impacts. Prescribed fire could be used to conserve grassland habitat but often has limited efficacy because many woody plants resprout after fire and rapidly reestablish abundance. If fire‐induced mortality could be increased, prescribed fire would be a more effective management tool. In California's central coast, shrub encroachment, especially of Baccharis pilularis (coyote brush), is converting coastal prairie into shrub‐dominated communities, with a consequent loss of native herbaceous species and open grassland habitat. B. pilularis has not been successfully controlled with single prescribed fire events because the shrub resprouts and reestablishes cover within a few years. We investigated whether two consecutive annual burns would control B. pilularis by killing resprouting shrubs, without reducing native herbaceous species or encouraging invasive plants. As expected, resprouting did occur; however, 2 years after the second burn, B. pilularis cover on burned plots was only 41% of the cover on unburned plots. Mortality of B. pilularis more than doubled following the second burn, likely maintaining a reduction in B. pilularis cover for longer than a single burn would have. Three native coastal prairie perennial grasses did not appear to be adversely affected by the two burns, nor did the burns result in increased cover of invasive species. Managers wanting to restore coastal prairie following B. pilularis encroachment should consider two consecutive annual burns, especially if moderate fire intensity is achievable.     

Invading Monotypic Stands of Phalaris arundinacea: A Test of Fire, Herbicide, and Woody and Herbaceous Native Plant Groups

Phalaris arundinacea L. is an aggressive species that can dominate wetlands by producing monotypic stands that suppress native vegetation. In this study invasion windows were created for native species in monotypic stands of P. arundinacea with either fire or herbicide. Three native species groups, herbaceous plants, herbaceous seeds, and woody shrubs, were planted into plots burned or treated with herbicide in the early spring. Fire did not create an effective invasion window for native species; there was no difference in P. arundinacea root and shoot biomass or cover between burned and control plots (p≥ 0.998). Herbicide treatment created an invasion window for native species by reducing P. arundinacea root and shoot biomass for two growing seasons, but that invasion window was fast closing by the end of the second growing season because P. arundinacea shoot biomass had nearly reached the shoot biomass levels in the control plots (p= 0.053). Transplant mortality, frost, and animal herbivory prevented the herbaceous species and woody seedlings from becoming fully established in the plots treated with herbicide during the first year of the experiment. Transplanted monocots had a greater survival than dicots. By the second growing season the herbaceous group had the greatest mean areal cover (5%), compared to the woody seedlings (3%) and seed group (0%). Long‐term monitoring of the plots will determine whether the herbaceous transplants will compete effectively with P. arundinacea and whether the woody species will survive, shade the P. arundinacea, and accelerate forest succession.     

Comparison of the regeneration after burning,cutting and ploughing in a Cistus ladanifer shrubland

The recovery of a Cistus shrubland after experimental treatments (cutting, burning and ploughing) was studied to determine qualitative and quantitative differences in the secondary succession. The study was carried out in three experimental plots of 100 m² each, in which five 1 m² sampling units were fixed and surveyed for five years, estimating cover percentage of each plant species. The data were analysed by means of affinity indexes (Sorensen and Steinhaus Indexes). The results indicated that the cover of the woody species increased progressively, but the degree of recovery depended on the treatment, as well as on the time afterwards. The number of these species remained almost constant throughout the whole period. The herbaceous species diversity was higher in the two first years in all treatments. When treatments were compared, the greatest diversity occurred after cutting. The regeneration of the community occurred by autosuccession in which the diversity decreased when the dominance of Cistus increased, unlike the general trend of the classical theories. The recovery was very quick after all three treatments, which can be interpreted as a great resilience of this ecosystem.     

Variability in seed germination at the interpopulation,intrapopulation and intraindividual levels of the shrub Erica australis in response to fire-related cues

In Mediterranean ecosystems, as well as in other fire prone ones, seedling establishment by some species is particularly favorable right after fire. It has been well established that many plants from these ecosystems have enhanced germination after exposing their seeds to heat or to chemicals related to the passage of fire. Less known it is how variable is this germination response among seeds from different sources (populations, individuals within a population, or parts of the plant within an individual) and whether such differences persist after exposing the seeds to germination-triggering, fire-related cues. In this work we studied the germination response to several fire-related factors of the Mediterranean, lignotuberous shrub Erica australis. This species produces a very combustible fuel, and resprouts vigorously after fire, but conditions for seedling establishment are poorly known. The objective of this study was to evaluate how variable was the germination response of this species among seeds from different sources, and whether this variability changed after exposing the seeds to fire-related factors. Seeds from three different sites, from different individuals at each site, and from different branches within each individual were set to germinate under control conditions or after exposing them to heat or to additions of nitrogenous compounds. Germination was highly variable among populations, individuals within populations or branches within individuals. Exposure to moderate heat promoted germination, but high temperatures were lethal. Differences at the various provenance levels were, however, also observed after heat exposure. Germination of seeds exposed to nitrogenous compounds increased, but differences among populations were always significant. These results indicate that the germination of Erica australis may be linked to disturbances, but it does not seem to be particularly related to passage of fire. On the contrary, this type of response appears more common in plants from heterogeneously disturbed environments. This revised version was published online in August 2006 with corrections to the Cover Date.     

Impact of habitat disturbance in the wetland forests of East Usambara,Tanzania

We evaluated habitat characteristics of East Usambara wetland forests. The abundance and species composition in the tree, shrub and herbaceous layers were enumerated in two sets of nested plots, one set in a natural wetland forest and the second in a wetland forest that had been disturbed by small‐scale gold mining activities. Each set had thirty‐six 1 m × 1 m plots for herbs, inside nine 5 m × 5 m plots for shrubs, in three 20 m × 20 m plots for trees. The habitat profile of herbaceous – shrub – tree layers was found to be sharply different from one obtained in previous studies at the surrounding nonwetland forests as were species composition and abundance. Unlike in the nonwetland forests, the herbaceous layer was thick, the shrub layer very thin and the woody species density and richness much lower. Disturbance significantly reduced woody cover and changed species composition in the herbaceous layer. Recovery of the woody vegetation was low. Wetland forests in the East Usambaras form a small fraction of the total area, but they are a unique biodiversity repository, they appear to be an important carbon dioxide sink and to reserve and purify water. They are sensitive to disturbance and need protection.