Long-term understory recovery after mechanical fuel reduction in Mediterranean cork oak forests

In the Euro-Mediterranean region, mechanical fuel reduction is increasingly used in response to the mounting occurrence of catastrophic wildfires, yet their long-term ecological effects are poorly understood. Although Mediterranean vegetation is resilient to a range of disturbances, it is possible that widespread fuel management at short intervals may threaten forest structural complexity and the persistence of some plant species and functional types, with overall negative consequences for biodiversity. We used a chronosequence approach to infer woody vegetation changes in the first 70 years after understory clearing in upland cork oak (Quercus suber) forests, and to assess how these are affected by treatment frequency. Across the chronosequence there was a shift between plant communities with contrasting composition, structure and functional organization. Understory cover increased quickly after disturbance and a community dominated by pioneer seeder and dry-fruited shrubs (Cistus ladanifer, C. populifolius, Genista triacanthos, and Lavandula stoechas) developed during about 15 years, but this was slowly replaced by a community dominated by resprouters and fleshy-fruited species (Arbutus unedo, Erica arborea) >40 years after disturbance. During the first 15 years there were rapid increases in woody species richness, vertical structural diversity, cover by Q. suber juveniles and saplings, and shrub cover at <1.5 m strata, which levelled off or slightly declined thereafter. In contrast, tree species richness, tree density and density of arboreal A. unedo and E. arborea, vertical structural evenness, and cover at >1.5 m strata increased slowly for >50 years. Treatment frequency showed strongly negative relationships with species richness, structural diversity and evenness, and horizontal and vertical understory cover, particularly that of slowly recovering species. These findings suggest that fuel reduction programs involving widespread and recurrent understory clearing may lead to the elimination at the landscape scale of stands with complex multi-layered understory occupied by large resprouters and fleshy-fruited species, which take a long time to recover after disturbance. Fuel management programs thus need to balance the dual goals of fire hazard reduction and biodiversity conservation, recognizing the value of stands untreated for >50 years to retain ecological heterogeneity in Mediterranean forest landscapes.     

Facilitation by two exotic Acacia: Acacia auriculiformis and Acacia mangium as nurse plants in South China

Because of their high growth rate and tolerance to bare soil, two exotic Acacia species, Acacia auriculiformis and Acacia mangium, have been commonly planted in degraded areas of South China. With their large canopies and ability to fix nitrogen, the two Acacia species have also been considered to act as nurse plants for understory plants. The current study clarified the nursing effects of the Acacia species by comparing microclimate characteristics and physiological traits of native plant seedlings at three sites: under the canopies of the each Acacia species and on bare land (open site). Although the sites were not replicated, the results indicated that adult trees of both Acacia species can facilitate native species, but that A. mangium has greater facilitating effects due to greater temperature buffering, radiation reduction, and nutrient amelioration. In response to facilitation, three species (Castanopsis hystrix, Michelia macclurei, and Manglietia glauca) with different shade-tolerant traits growing under Acacia canopies expressed distinct adaptations. For the three species, the chlorophyll fluorescence curves of rETR and ΔF/Fm′ were higher under A. auriculiformis and on the open site than under A. mangium. The maximum quantum yield in PSII(Fv/Fm) in diurnal changes of the three species showed that all the Fv/Fm values were between 0.70 and 0.84 and that the Fv/Fm values were mostly higher under A. mangium than on the open site or under A. auriculiformis. Total chlorophyll content and both chlorophyll a and b contents in the three species were higher under the Acacia species than on the open site, while chlorophyll a/b ratio was higher on the open site. In contrast, the carotenoid content in C. hystrix and M. macclurei was lower under the two Acacia species than on the open site, while the opposite was true for M. glauca. The results demonstrate that the adaptation of the understory species to abiotic environmental factors is not restricted to a single mechanism but apparently involves a group of interrelated, adaptive suites. And also these adaptations were species-specific and especially related to their shade tolerance.     

Sphagnum community change after partial harvest in black spruce boreal forests

In eastern Canada, boreal forests develop structural diversity in association with time since stand replacing fire. In some regions, this is associated with significant changes in the bryophyte community (Sphagnum moss invasion) and paludification (thick waterlogged forest floor development). The bryophyte community responds to opening of the canopy, and increasing moisture by replacement of slow growing species by faster growing Sphagnum spp. (e.g. magellanicum, fallax) that are dependent on constant hydration. Within a forest management context, partial harvest systems have been proposed as a strategy to maintain structural diversity, which is currently not accomplished with low retention systems. However, it is unknown whether these interventions will effectively accelerate community succession. The questions addressed in this study were: (1) is the composition of Sphagnum colonies in partially cut stands more similar to old-growth communities than in control, and low retention cut stands, (2) what aspects of harvest disturbance drive these changes, and (3) is the growth rate of Sphagnum capillifolium (an early successional shade tolerant species) different in partial versus low retention harvest systems? After harvest, Sphagnum patch size was reduced by 19.8% and 11.7% after low retention and partial harvest, respectively. While trends were not constant across three separate partial cut trials, the proportion of Sphagnum magellanicum, Sphagnum fallax and Sphagnum fuscum increased compared to controls and low retention 1–2 years after harvest. Models of percent Sphagnum cover indicated machinery track cover, percent cover of vascular plants, and patch depth were positive factors, while the influence of open canopy varied among species. Despite the inclusion of individual disturbance variables, the summary variable ‘treatment’ was significant in all models. Growth of S. capillifolium in partial cuts was intermediate to growth rates in control and low-retention cuts. Growth was positively influenced by slash cover and, contrary to the patch level, negatively influenced by track cover. These results indicate that partial harvest does represent an intermediate level of disturbance, as direct and indirect harvest effects were reduced, as was Sphagnum death. Change in composition 1 and 2 years after harvest indicates that partial harvests may effectively shift the bryophyte community towards an older community type and may thus be used to create landscape diversity. Long term trends and entire community compositions need to be assessed before this can be stated definitively. However, as paludified stands are less productive, the capacity of these partially harvested sites to produce merchantable timber is questioned.     

Quercus calliprinos regrowth advantage under grazing in Mediterranean maquis and its management implications

The post disturbance natural succession of the Mediterranean maquis, which turns open and species-rich landscapes into dense, closed stands of sclerophyllous woody vegetation is a principal threat to plant and animal diversity. Therefore, in the absence of traditional agricultural disturbance, active management regimes that include cutting and grazing are proposed to preserve biodiversity. The Mediterranean woody vegetation in Israel is strongly dominated by one species - the evergreen sclerophyllous Quercus calliprinos (Kermes oak). We hypothesized that under cutting and grazing, the evergreen Q. calliprinos has a relative regrowth advantage over other competing tree species. Here we examined the effect of grazing and the effect of tree structural traits on the regrowth after clear cutting of all trees in our study plots at Mt. Meron LTER site, Israel. All trees were removed from five blocks of 2000 m² and each block was divided into two plots, five of which were exposed to grazing livestock while five were wire-fenced and ungrazed. The regrowth rate of Q. calliprinos under grazing was higher than that of all other tree species suggesting that in the long-term, under such a conservation management regime, the dominance of the evergreen sclerophyllous Q. calliprinos over the deciduous tree species will increase and consequently will decrease plant and animal diversity. Therefore, we conclude that to protect landscape and species diversity in Mediterranean ecosystems dominated by evergreen oaks, when cutting and grazing are applied, special care must be paid to trees that are more negatively affected by such treatment.     

The effect of herbivory on seedling survival of the invasive exotic species Pinus radiata and Eucalyptus globulus in a Mediterranean ecosystem of Central Chile

Herbivory may be an important factor affecting seedling survival of exotic species invading new habitats. We evaluated the effect of vertebrate herbivory on the seedling survival of two widely planted and invasive tree species (Pinus radiata and Eucalyptus globulus), in a Mediterranean-type ecosystem of central Chile. An important role of herbivory on seedling survival of these two species in their introduced ranges has previously been documented. However, this has mainly been evaluated in forest plantations where habitat and vegetation conditions differ from wild habitats in which invasion occurs. We planted seedlings with and without protection against vertebrate herbivores in different aspects (a mesic south-facing slope and a xeric north-facing slope) and vegetation cover (open sites and sites with patchy tree cover). We found that regardless of aspect or vegetation cover, herbivory, in this case mainly caused by exotic vertebrates, significantly and negatively affected seedling survival of both species. However, while the effect of herbivory on P. radiata was significant in every vegetation and habitat condition, for E. globulus, the effect of herbivory was significant only for open sites in the mesic habitat. These results suggest that, as observed in forestry plantations, vertebrate herbivory may constrain seedling establishment of these two exotic trees and potentially impede the invasion. However, the importance of herbivory in controlling exotic species may vary depending on the vegetation and habitat conditions in some species such as E. globulus.     

Fungal community succession following wildfire in a Mediterranean vegetation type dominated by Pinus pinaster in Northwest Spain

This study examined the succession of fungal communities following fire in a Mediterranean ecosystem dominated by Pinus pinaster Ait. in northwestern Spain. A large wildfire occurred in August 2002. During the autumn seasons from 2003 to 2006, fruiting bodies were collected and identified, production in burned (early stage) and unburned (late stage) areas was measured. For statistical analysis, data were grouped into the following four categories: saprotrophic, mycorrhizal; edible and inedible. A total of 115 fungal taxa were collected during the four sampling periods (85 in the late and 60 in the early stage). The number of mycorrhizal species increased from early to late succession and there were shifts in community composition. After fire, pyrophytic species such as Pholiota carbonaria, Peziza violacea, Rhizopogon luteolus and Rhizopogon sp. appeared. Fire strongly affected the production of fungal species in the studied area. Thus, yields in the early stage treatment were significantly lower than those observed in the late stage. Total fungal fresh weight decreased from 209.95 kg fw ha⁻¹ in the late stage to 162.45 kg fw ha⁻¹in the early stage when richness and production of mycorrhizal species and production of edible fungi were significantly lower. Fresh weight for saprotrophic and inedible species was higher than for mycorrhizal fungi in the early stage treatment. The results obtained can be useful to forest managers for optimization of management and harvesting of these increasingly appreciated non-timber resources.     

Coarse woody debris in late-successional Picea abies forests in northern Europe: Variability in quantities and models of decay class dynamics

Coarse woody debris pools and their decay class dynamics were studied in three areas of unmanaged boreal forest in northern Finland, and in the Murmansk and Arkhangelsk provinces in northwestern Russia. The study areas had varying climatic and edaphic conditions, and disturbance histories. Living and dead trees (diameter at 1.3 m height ≥10 cm) were measured in five late-successional Picea abies-dominated stands in each of the three areas. Wood density and time since death were determined from randomly sampled dead P. abies, and their decay class dynamics were modeled using stage-based matrix models.     

Modeling the effects of alternative management strategies on forest carbon in the Nothofagus forests of Tierra del Fuego,Chile

The impact of forest management activities on the ability of forest ecosystems to sequester and store atmospheric carbon is of increasing scientific and social concern. The nature of these impacts varies among forest ecosystems, and spatially and temporally explicit ecosystem models are useful for quantifying the impacts of a number of alternative management regimes for the same forest landscape. The LANDIS-II forest dynamics simulation model is used to quantify changes to the live overstory and coarse woody debris pools under several forest management scenarios in a high-latitude South American forest landscape dominated by two species of southern beech, Nothofagus betuloides and N. pumilio. Both harvest type (clearcutting vs. partial overstory retention) and rotation length (100 years vs. 200 years) were significant predictors of carbon storage in the simulation models. The prompt regeneration of harvest units greatly enhanced carbon storage in clearcutting scenarios. The woody debris pool was particularly sensitive to both harvest type and rotation length, with large decreases noted under short rotation clearcutting. The roles of extended rotations and partial overstory retention are noted for enhancing net carbon storage on the forest landscape.     

Patterns of spread of Pinus contorta Dougl. ex Loud. invasion in a Natural Reserve in southern South America

Pinus species, which have formed the foundation of commercial forestry industry in many countries, are known to be invasive in natural ecosystems, especially in the Southern Hemisphere. Pinus contorta is considered one of the most aggressively invasive plantation species. In this paper we aim to: (a) determine patterns of P. contorta invasion in relation to its size and age structure and spatial distribution and (b) determine the effect of vegetation cover on its regeneration. For this purpose, we have chosen the Malalcahuello National Reserve in south-central Chile, which is dominated by Araucaria araucana forests. In the three P. contorta trial plots, attributes were measured in order to describe its current structure. We selected one of the three P. contorta trial plots and set eight 50 m wide and 125 m centrifugal transects starting at the North Azimuth (0°) orientation and then one each 45°. In each transect 25 circular plots of 2.5 m radius were established every 25 m. In each plot, we measured collar diameter (CD) for each P. contorta individual and adjusted a diameter–age function. We recorded the presence of cones for each individual P. contorta within the plots and the total number of P. contorta individuals. In each plot, we measured percent vegetation cover (grass, understory and canopy). The selected parent stand showed a decrease in density from the original plantation spacing, from 2500 to 150 plants ha⁻¹. Of all sampled individuals, only a 38% had cones. We found reproductive structures in trees as young as 5-year-old. Regeneration was found in all transects. Mean density for the area was 1600 plants ha⁻¹, and the greater plant number was found in the West transect, reaching an average of 6600 plants ha⁻¹. Through the interpolation performed with the kriging method, a map of the area with the spatial gradient of plant density was obtained. Naturally regenerated individuals of P. contorta occupied an area of 78 ha where the most distant individual is located in the southeast direction at an average distance of 1200 m from the three study plots. We determined that a positive association exists between P. contorta and the species A. araucana and Nothofagus antarctica. The capacity for early and consistent reproduction and the establishment of individuals dispersed at great distances from the original plots allow us to conclude that P. contorta has great potential as an invasive species in forests of this area of the Andes of southern South America. Invasion of P. contorta has many important implications for the conservation of native forests in our region including diminished regeneration of A. araucana.