Alternative silvicultural practices with variable retention to improve understory plant diversity conservation in southern Patagonian forests

Understory plants could can act as indicators of temperate forest sustainability, health and conservation status due to their importance in ecosystem function. Harvesting impacts on understory plant diversity depends on their intensity. Variable retention has been proposed to mitigate the harmful effects of timber harvesting, but its effectiveness remains unknown in southern Patagonian Nothofagus pumilio forests. The objectives of this study were to: (i) define a baseline of understory plant diversity in old-growth forests along a site quality gradient and under canopy gaps; (ii) evaluate stands with three different variable retention treatments compared to old-growth forests; and (iii) assess temporal changes during 4 years after harvesting (YAH). A 61 ha N. pumilio forest was selected. Understory plant (Dicotyledonae, Monocotyledonae and Pteridophyta) richness, cover (including woody debris and bare forest floor) and aboveground dry biomass were characterized in summer for 5 years. Before harvesting, baseline samples were conducted along a site quality gradient and outside/inside canopy gaps. Analyzed treatments include a control of old-growth forest (OGF) and three different harvesting treatments with variable retention: (i) dispersed retention (DR) of 30 m² ha⁻¹ (20-30% retention); (ii) aggregated retention (AR) with one aggregate per hectare and clear-cuts (28% retention); and (iii) combined dispersed and aggregated retention (DAR) with one aggregate per hectare and dispersed retention of 10-15 m² ha⁻¹ (40-50% retention). Data analyses included parametric and permutational ANOVAs, multivariate classification and ordinations.Before harvesting, 31 plant species were found, where richness, cover and biomass were directly related to site quality. The presence of canopy gaps did not have a significant impact on the measured variables. After harvesting, 20 new species appeared from adjacent associated environments (two from N. antarctica forests and 18 from grasslands and peatlands). At the stand level, understory values were higher in AR > DR > DAR > OGF. Most (81-95%) plant richness at baseline conditions was conserved in all treatments, where inside the aggregates understory remained similar to OGF. Combination of aggregated and dispersed retention (DAR) better limited exotic species introduction and protected sensitive species, improving conservation in harvested stands. Changes in understory variables were observed after the first YAH in all treatments; greater changes were observed in the harvested areas than in aggregates. Changes stabilized at the fourth YAH. As a conclusion, the location of retention aggregates should be selected to preserve species understory diversity of more speciose and diverse habitats or particularly uncommon stands. Implementation of different kinds (patterns and levels) of retention for improvement of biodiversity conservation in harvested forests should be included in timber and forest management planning.     

Performance of three-layer composites with densified surface layers of Nothofagus pumilio and N. antarctica from Southern Patagonian forests

Nothofagus pumilio and N. antarctica forests of Southern Patagonia, Argentina are currently being managed for production of saw logs with fast growing conditions. The result of these management strategies is faster growing forests, but also an increase in the percentage of low-density wood. The motivation for this study was to find a way to valorise this low-density wood. Surface and bulk densification treatments were applied these wood species and were then applied as face layers in three-layer composites. The mechanical properties of three-layer composites were studied. The modulus of elasticity, modulus of rupture, and modulus of hardness were improved compared to many types of structural composite lumber. This increase in mechanical properties of N. pumilio and N. antarctica wood opens the possibility for its use in structural composites and added value to these otherwise underutilised and undervalued species.     

Foliar anatomical and morphological variation in Nothofagus pumilio seedlings under controlled irradiance and soil moisture levels

Foliar anatomy and morphology are strongly related to physiological performance; therefore, phenotypic plasticity in leaves to variations in environmental conditions, such as irradiance and soil moisture availability, can be related to growth rate and survivorship, mainly during critical growth phases, such as establishment. The aim of this work was to analyze changes in the foliar internal anatomy (tissue proportions and cell dimensions) and external morphology (leaf length, width and area) of Nothofagus pumilio (Poepp. et Endl.) Krasser seedlings growing in a greenhouse under controlled irradiance (three levels) and soil moisture (two levels) during one growing season (measured three times), and to relate them to physiological traits. Three irradiance levels (4, 26 and 64% of the natural incident light) and two soil moisture levels (40 and 80% soil capacity) were evaluated during November, January and March. Internal foliar anatomy of seedlings was analyzed using digital photographs of histological cuttings, while leaf gross morphology was measured using digital calipers and image analysis software. Most internal anatomical variables presented significant differences under different irradiance levels during the growing season, but differences were not detected between soil moisture levels. Palisade parenchyma was the tissue most sensitive to irradiance levels, and high irradiance levels (64% natural incident light) produced greater values in most of the internal anatomical variables than lower irradiance levels (4-24% natural incident light). Complementarily, larger leaves were observed in medium and low irradiance levels, as well as under low soil moisture levels (40% soil capacity). The relationship of main results with some eco-physiological traits was discussed. Foliar internal anatomical and external morphological plasticity allows quick acclimation of seedlings to environmental changes (e.g., during harvesting). These results can be used to propose new forest practices that consider soil moisture and light availability changes to maintain high physiological performance of seedlings.     

Alternative silvicultural practices with variable retention improve bird conservation in managed South Patagonian forests

Alternative silvicultural approaches to timber management, such as regeneration treatments with different degrees of stand retention, may mitigate negative effects of clear-cutting or shelterwood cuts in forested ecosystems, including changes in old-growth forest bird communities. The aims of this work were: (a) to compare bird species richness and densities among different silvicultural designs with variable retention (dispersed and/or aggregated) and unmanaged primary forests, and (b) to assess temporal changes at community and species levels before and after treatments. A baseline avian survey was conducted prior to harvesting to evaluate canopy gap presence and forest stand site quality influences. Subsequent to harvesting, data on bird species richness and density were collected by point-count sampling during the summer season for 5 consecutive years (4 treatments × 5 years × 6 sampling points × 5 counts). Bird species richness and density (15 species and 9.2 individuals ha⁻¹) did not change significantly with forest site quality of the stands and canopy gap presence in unmanaged forests. However, both variables were significantly modified in managed forests, increasing over time to 18 species and reaching to 39 individuals ha⁻¹. Inside the aggregated retention, bird communities were more similar to unmanaged primary forests than those observed within the dispersed retention or in clear-cuts. Opting for a regeneration method with dispersed and aggregated retention has great potential for managing birds in Nothofagus pumilio forests. This method retained enough vegetation structure in a stand to permit the establishment of early successional birds (at least in dispersed retention), and to maintain the bird species of old-growth forests which could persisted in the retention aggregates.     

Differential forage use between large native and domestic herbivores in Southern Patagonian Nothofagus forests

Plant—animal interactions at the landscape level become particularly relevant when land use is diversified. Nothofagus forests in southern Patagonia have been used for timber and cattle grazing purposes during the last century, causing livestock to increase (Bos taurus and Ovis aries) and large native herbivores, such as guanaco (Lama guanicoe), either maintained or decreased their populations. Within this scenario, feeding interactions between guanaco and domestic herbivores were analysed in a mosaic of open habitats and different Nothofagus forest types and management histories, whereby a total of six habitat types were identified through satellite image analysis. A total of 205 floristic surveys were conducted to characterize the plant species composition at the landscape level. Diet composition of herbivores was assessed once each season during a year, using microhistological analysis of feces. Results showed higher plant richness in open lands and lower in closed, unmanaged forests. Overall, 43 plant taxa were detected in herbivore feces, which represent 56% of plant richness detected in the field. Both guanacos and domestic herbivores included mainly grasses in their diet, which were found predominantly in open lands. Tree seedlings and saplings were consumed by all herbivores, as well, except during winter. Differential forage use between guanaco and domestic herbivores in southern Patagonian Nothofagus forests did exist. However, competition for available resources among theses herbivores resulted in an alternation of feeding sites, which varied throughout the year. Management plans in southern Southern Patagonia (livestock, silvopastoral plans and timber harvesting) do not consider the direct or indirect consequences on guanaco populations. The challenge, therefore, is to generate management decisions to avoid either guanacos or domestic herbivores from becoming detrimental to the sustainability of managed forested ecosystems.     

Silvopastoral use of Nothofagus antarctica in Southern Patagonian forests, influence over net nitrogen soil mineralization

In most temperate forest, nitrogen (N) is considered a limiting factor. This becomes important in extreme environments, as Nothofagus antarctica forests, where the antecedents are scarce. Thinning practices in N. antarctica forests for silvopastoral uses may modify the soil N dynamics. Therefore, the objective of this work was to evaluate the temporal variation of soil N in these ecosystems. The mineral extractable soil N, net nitrification and net N mineralization were evaluated under different crown cover and two site quality stands. The mineral N extractable (NH₄ ⁺–N + NO₃ ^?–N) was measured periodically. Net nitrification and net N mineralization were estimated through the technique of incubation of intact samples with tubes. The total mineral extractable N concentration varied between crown cover and dates, with no differences among site classes. The lowest and highest values were found in the minimal and intermediate crown cover, respectively. In the higher site quality stand, the annual net N mineralization was lower in the minimal crown cover reaching 11 kg N ha^?1 year^?1, and higher in the maximal crown cover (54 kg N ha^?1 year^?1). In the lower site quality stand there was no differences among crown cover. The same pattern was found for net nitrification. Thinning practices for silvopastoral use of these forests, keeping intermediate crown cover values, did not affect both N mineralization and nitrification. However, the results suggest that total trees removal from the ecosystem may decrease N mineralization and nitrification.     

Regeneration of Fitzroya cupressoides after indigenous and non-indigenous timber harvesting in southern Chilean forests

Fitzroya cupressoides (Cupressaceae) is an endemic and long-lived conifer of southern Chile and Argentina (40–43° S). This species has been subject to continuous exploitation since the 16th century, causing extensive population decline. Historically, the main labour force for the exploitation of F. cupressoides (alerce) was the indigenous Mapuche-Huilliche population, first under the command of the Spanish settlers and later, of non-indigenous Chileans. In coastal forests, timber of alerce has been harvested by Huilliche communities as well as by Chilean and international forestry companies. Records of the regeneration of this pioneer tree after exploitation in the Andean mountains have generally shown limited regeneration depending on the intensity of harvest. Because indigenous exploitation does not use machinery for timber harvesting, and is supposedly less utilitarian than commercial harvest, I propose that areas in the Coastal Range harvested by Huilliche communities should present higher regeneration of alerce than areas harvested by forestry companies. To test this hypothesis, I sampled 10 stands harvested and abandoned by forestry companies and 10 stands harvested by Huilliche communities in the coastal range of the Osorno Province (41° S, 400–800 m). In each stand, I estimated the density of regeneration (sapling stage), number of stumps, number of live adults, and the number of standing and fallen dead alerce. Each stand was characterized by elevation, forest-type, incidence of fire, and vegetation cover. Results showed that forests harvested by Huilliches had higher numbers of live, remnant adult trees with a dbh ≥ 60 cm than forests harvested by timber companies. The number of stumps with a dbh ≥60 cm was significantly higher in stands harvested by timber companies than in Huilliche stands. Despite large differences in sapling densities among stands, regeneration density of alerce was unrelated to the type of harvest used by indigenous people or forestry companies.     

Spores of ectomycorrhizal fungi as inoculants for Nothofagus pumilio and exotic conifer seedlings in Patagonia, Argentina: their activity and conservation

Ectomycorrhizal (EM) fungi incorporation in nursery production is necessary for afforestation with exotic tree species and native forests restoration programs. The fact that spore use as EM inocula is inexpensive and effective, makes it an option more realistic than vegetative inoculum production for many regions around the world. To determine spore activity from EM species selected as inoculants for Nothofagus pumilio and exotic conifers planted in Patagonia, two conservation methods were applied (i.e., dried fruit bodies and spore slurries), and their change over time was assessed (over 8–9 months). Spore activity decreased significantly with time for both native and exotic EM species. Conservation methods showed no significant differences for N. pumilio EM species considered together or for exotic conifer EM species. However, spore activity of different EM species behaved differently with conservation method and over time when considered separately. Taxa which better kept spore activity over time were Austropaxillus statuum and Setchelliogaster fragilis for N. pumilio, and Inocybe kauffmanii for exotic conifers. However, considering together fruit body spore density and spore activity, the species Hallingea purpurea and Cortinarius sp. also appeared as suitable for N. pumilio and Rhizopogon roseolus, R. villosulus, Suillus luteus and S. lakei for exotic conifers. Spore density found in EM fungal fructifications were also established in order to estimate fructification weights necessary to apply a reference dose of active spores in inoculation programs.     

Regeneration of Populus nine years after variable retention harvest in boreal mixedwood forests

Aspen and balsam poplar regeneration from root suckers were assessed in boreal mixedwood forests nine years after logging in a variable retention experiment (EMEND Project—Ecosystem Management Emulating Natural Disturbance) located north of Peace River, Alberta, Canada. Five levels of retention of mature trees (2%, 10%, 20%, 50% or 75% of the original basal area) were applied in stands dominated by aspen, white spruce or mixtures of the two species. Basal area of aspen (or that of aspen plus balsam poplar combined) prior to logging strongly influenced sucker density of aspen (or aspen + balsam poplar combined) and in some cases their growth. Nine years after harvest there was a decline in sucker density and volume ha⁻¹ with increasing retention levels of aspen (or both poplars combined); sucker density declined by 50% when only 20% of the original basal area was left in the stand. Retaining mature spruce trees in the stand had little influence on the number of suckers but did affect their total volume ha⁻¹. Thus, we suggest that by knowing stand aspen and balsam poplar density prior to logging and varying levels of retention of aspen and balsam poplar or conifers at harvest, the density of Populus regeneration can be predicted by managers, thereby allowing them to create a range of mixedwood conditions.     

Container seedlings outperform barefoot stock: Survival and growth after 10 years

Ponderosa pine, Jeffrey pine, and Douglas-fir seedlings were planted in container or bareroot form at three elevations in northern California. At the lowest elevation (762 m), container seedlings of ponderosa pine were significantly taller than bareroot seedlings at ages 4 and 10, and had breast-height diameters that were significantly larger than bareroot counterparts at age 10. Survival of Douglas-fir container seedlings was significantly greater than that of barefoot seedlings for all ages tested. At the mid-elevation site (1220 m), container seedlings of ponderosa pine and Douglas-fir were significantly larger in breast-height diameter than bareroot seedlings at age 10. Douglas-fir container seedlings survived significantly better at all ages than barefoot seedlings. At the highest elevation (1662 m), seedling height and diameter did not differ significantly, but survival of container seedlings was significantly higher than barefoot seedlings for both pine species at all ages tested.     

Carbon isotope discrimination and water stress in trembling aspen following variable retention harvesting

Variable retention harvesting (VRH) has been proposed as a silvicultural practice to maintain biodiversity and ecosystem integrity. No previous study has examined tree carbon isotope discrimination to provide insights into water stress that could lead to dieback and mortality of trees following VRH. We measured and compared the carbon isotope ratios (delta(13)C) in stem wood of trembling aspen (Populus tremuloides Michx.) before and after VRH. Eight trees were sampled from isolated residual, edge and control (interior of unharvested stand) positions from each of seven plots in three regions (Calling Lake and Drayton Valley, Alberta and Lac Duparquet, Québec). After VRH, the general trend in mean delta(13)C was residual > edge > control trees. Although this trend is indicative of water stress in residual trees, it also suggests that edge trees received some sheltering effect, reducing their stress compared with that of residuals. A strong inverse relationship was found between the delta(13)C values and the mean annual precipitation in each region. The trend in mean delta(13)C signature was Calling Lake > Drayton Valley > Lac Duparquet trees. These results suggest that residual or edge trees in drier regions are more likely to suffer water stress following VRH. We also observed a trend of greater delta(13)C in stout trees compared with slender trees, both before and after VRH. The evidence of greater water stress in stout trees likely occurred because of a positive relationship between stem diameter and crown volume per basal area. Our results provide evidence that water stress could be the driving mechanism leading to dieback and mortality of residual trees shortly after VRH. Additionally, the results from edge trees indicate that leaving hardwood residuals in larger patches or more sheltered landscape positions could reduce the water stress to which these trees are subjected, thereby reducing dieback and mortality.     

Survival of southern pine seedlings after inoculations with Pythium and cold storage in the presence of peat moss

Cold storing bareroot pine (Pinus spp.) seedlings grown in the southern U.S. for as little as 1 week in a cooler (just above freezing) in the fall (November to mid‐December) has been shown to reduce seedling survival after outplanting. In contrast, survival of container‐grown seedling is typically not affected when stored for 4 weeks in coolers in November and December. Wounds sustained by seedlings as they are lifted from nursery beds may allow Pythium spp. to infect bareroot seedling roots. Once in the cool, moist storage environment, Pythium multiplies and may result in seedling mortality after outplanting. Bareroot loblolly pine (Pinus taeda) and container‐grown loblolly, longleaf (Pinus palustris), slash (Pinus elliottii) and shortleaf pine (Pinus echinata) seedlings were inoculated with either Pythium dimorphum or Pythium irregulare, cold stored with or without peat moss and monitored for survival after outplanting. Peat moss did not increase bareroot loblolly pine survival or reduce Pythium populations when seedlings were inoculated with Pythium prior to storage. Pythium irregulare reduced survival of longleaf and shortleaf pine grown in peat moss and perlite, respectively. Pythium did not affect loblolly or slash pine, but wounding their roots did reduce seedling survival when grown in containers.     

Survival,early growth and impact of damage by late-spring frost and winter desiccation on Douglas-fir seedlings in southern Sweden

Introduction of non-native species, such as Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), can be a means of mitigating the effects of climate change by meeting the growing demand for biomass and high quality wood. The aim of this study was to investigate early growth, survival and damage from late-spring frost and winter desiccation. A provenance trial with four coastal and three interior provenances of Douglas-fir originating from British Columbia, Canada, was established in Southwest Sweden (56°43′N, 13°08′E). Seedling height, length of the leading shoot, and occurrence of frost damage, were measured after one, three, and six growing seasons. Timing of bud break in spring was also observed. The interior Douglas-fir were more frequently damaged by late-spring frost compared to the coastal Douglas-fir. The interior Douglas-fir still had a higher survival after six growing seasons compared to the coastal variety. All provenances were damaged by winter desiccation, but the provenances originating from the coastal area were more severely damaged. Choice of variety may reduce the risk for either late-spring frost or winter desiccation.     

Refugial forests of the southern Appalachians: photosynthesis and survival in current-year Abies fraseri seedlings

Fraser fir (Abies fraseri (Pursh) Poiret) is an endemic, high-elevation conifer confined to six relict mountaintop communities in the southern Appalachian Mountains, USA. High adult mortality has occurred over the past 50 years, possibly the result of an introduced insect (Adelges piceae Ratzeburg), air pollution, or both. Knowledge of the mechanisms of and limitations to seedling establishment may allow reestablisment and perpetuation of this unique community type, notwithstanding global climate change. We monitored seedling emergence and mortality in relation to photosynthetic performance and water relations in microsites differing in canopy openness (sunlight exposure) over the summer of 2004. Abundance of cotyledonous seedlings in early summer was 2.3 times greater (849 versus 366 seedlings m(-2)) in microsites with lower sky exposure (greater canopy closure) than in microsites with greater sky exposure (greater canopy openness). In contrast, late-season abundance and survival were greater in areas beneath more open canopies than in areas beneath less open canopies (3.3 times and 11.7 times greater, respectively). However, newly emerged seedling survival in a completely open site (no overhead canopy) was zero, despite an initial density of 124 seedlings m(-2). Seedling water status was similar in open- and closed-canopy sites (-0.52 and -0.74 MPa, respectively). Photosynthetic carbon gain was higher in newly emerged seedlings at open canopy than at closed canopy sites, especially during early morning. Based on photosynthetic light response curves and measured sunlight regimes, seedlings in open canopy sites were estimated to assimilate 3.3-4.5 times more carbon than seedlings at closed sites. Reductions in carbon gain of closed-site seedlings, primarily a result of limited sunlight, corresponded to substantial increases in seedling mortality (98 versus 79% in open canopy sites). Thus, sunlight exposure, which reflects overstory canopy structure, appears to be an important factor influencing newly emerged seedling survival and distribution.     

Survival and growth dynamics of red spruce seedlings planted under different forest cover densities and types

New Forests - Advance regeneration of red spruce (Picea rubens Sarg.) is scarce in many forest stands, due to past clearcuts and heavy harvesting of large seed trees in partial cuts. Understory...     

Threshold effects of variable retention harvesting on understory plant communities in the boreal mixedwood forest

We studied the effects of six levels of dispersed green-tree retention (GTR) harvesting (clearcut (0%), 10%, 20%, 50%, and 75%, and unharvested reference (100%)) on understory plant communities in the 8th growing season post-harvest in the mixedwood boreal forest in northwestern Alberta. For the partial harvest treatments (10%, 20%, 50%, 75%) sample plots were located in the partially harvested (retention) strips as well as in the intervening machine corridors used by the harvesting equipment. The understory plant community was significantly influenced by the gradient of retention level. The cover of understory vegetation, especially graminoids, increased with increasing harvesting intensity for the retention strips and overall considering both plots types. Species richness was unaffected by retention level but did decrease as tree density increased. Lower levels of retention lead to increased abundance of early successional, shade-intolerant species. The results suggest a threshold in understory response to GTR harvesting between the 10% and 20% retention treatments. In terms of understory cover and composition, machine corridors within partially harvested forests resembled clearcuts. The results suggest that retaining more than 10% during GTR harvesting could have significant benefits in terms of maintaining understory plant communities more similar to unharvested reference forest.     

Timber management with variable retention in Nothofagus pumilio forests of Southern Patagonia

Forestry practices integrating ecological and social criteria have been replacing those based only on economic values. Traditional silviculture, such as shelterwood cuts (SC), transforms uneven-aged original stands to an even-aged managed forest. Recently, other methods have proposed to conserve some of the original heterogeneity of the old-growth forests. One proposal leaves 30% of the timber quality forest area as aggregated retention and 20% basal area as dispersed retention. The aim of this study was to analyze the feasibility of timber management with aggregated and dispersed retention in Nothofagus pumilio old-growth forests by analyzing timber and harvesting yield potential compared with traditional regeneration systems. Also, remnant tree stability of aggregated retention was analyzed.     

Mortality in relation to growth rate and soil resistance varies by species for underplanted Nothofagus seedlings in scarified shelterwoods

This study evaluated early mortality patterns of planted Nothofagus dombeyi (Mirb.) Blume (evergreen, shade intolerant) and Nothofagus alpina (P. et E.) Oerst. (deciduous, mid-tolerant to shade) seedlings established in scarified gaps in a Nothofagus old-growth forest in the Chilean Andes that was subjected to a shelterwood cut. Soils were scarified with heavy machinery that removed up to 40 cm of the upper horizons (e.g. it completely removed the litter layer and organic horizon, and in some cases partially the A horizon) to eliminate bamboo (Chusquea culeou E. Desv) shoots and rhizomes from the scarified areas. Two years after planting, seedlings were growing free of understory competition and mortality patterns were assessed using logistic regression models with the following predictor variables: (1) basal diameter growth in the prior year, and (2) soil strength measured as resistance to penetration. The probability of mortality of both species was highly predictable based on recent basal diameter growth. N. dombeyi seedlings appeared to be less affected by mortality, as reflected in lower sensitivity of mortality at lower growth rates compared to N. alpina. High values of resistance to penetration (>2,000 kPa) induced a higher probability of mortality in N. alpina (up to 40 % at 3,500 kPa). In contrast, N. dombeyi mortality was not significantly influenced by soil strength, which illustrates the more stress-resistant and pioneer character of this species. This study showed that recent growth rates can be used as a predictor of future mortality, but these relationships need to be calibrated for each species. In addition, our results suggest that species characteristics that relate to resource limitations, such as shade tolerance, are not adequate to understand effects of other stressors, such as soil disturbance, on mortality patterns.