Wind speed and crown class influence the height–diameter relationship of lodgepole pine: Nonlinear mixed effects modeling

Wind-induced mechanical perturbation is one of the mechanisms determining the height–diameter relationship of trees, but its effects have largely been neglected in past height–diameter models. In this study, we examined the effects of including wind speed as a regressor in the height–diameter model for lodgepole pine (Pinus contorta var. latifolia Engelm.). We also tested the hypothesis that the height–diameter relationship differs between trees of different crown classes (dominant and subordinate trees), and that a mixed effects height–diameter model developed at the crown class level should achieve a better prediction than that developed at the plot level. The results showed that including wind speed significantly improved the fit of the height–diameter model. The results also showed that the height–diameter relationships differ significantly between the two crown classes. The mixed effects model developed based on crown classes of individual trees had better fit with reduced residual variance and decreased Akaike's information criterion and the Schwarz's Bayesian information criterion than the plot-based mixed model. Evaluation of the developed model suggests the crown class-based height–diameter model has better prediction in terms of prediction error and precision. The developed crown class-based mixed effects model can be used to provide more accurate prediction of tree heights for lodgepole pine from their diameters.     

Evaluating the effect of drier and warmer conditions on water use by Quercus pyrenaica

Under climate change, severe and recurrent droughts can reduce forest production and cause widespread tree dieback. The response of different vegetation types to climate change can vary greatly and, therefore, must be individually assessed. This study was carried out in a Mediterranean oak forest (Quercus pyrenaica) subject to seasonal summer drought. To examine the response of the forest to the climate conditions predicted under climate change, a Soil–Vegetation–Atmosphere Transfer model [SPA, Williams, M., Rastetter, E.B., Fernandes, D.N., Goulden, M.L., Wofsy, S.C., Shaver, G.R., Melillo, J.M., Munger, J.W., Fan, S.M., Nadelhoffer, K.J. 1996. Modelling the soil-plant-atmosphere continuum in a Quercus–Acer stand at Harvard Forest: the regulation of stomatal conductance by light, nitrogen and soil/plant hydraulic properties. Plant, Cell, Environment 19, 911–927] was used. The model was parameterized using mostly local measurements (independent of the verification data) and tested against in situ sap flow measurements obtained during year 2007. The predictions of the model were broadly consistent with the observed dynamics of sap flow (the model explained 71% of the variance in daily transpiration and 75% of half-hourly sap flow), leaf water potentials and soil water content. Once the model had been validated, simulations were carried out under warmer and dryer conditions. Predicted warmer conditions (4 °C) caused a moderate increase in total simulated transpiration. Less frequent precipitation (40% longer dry periods between rainfall events) had very little effect on transpiration. In contrast, transpiration was reduced by 17% when the soil water reserves at the beginning of the summer were lower than in 2007, corresponding to those measured in a very dry year (2005). The reduction was exacerbated when changes in temperature and rainfall were also considered (up to 28% decline in transpiration). The higher atmospheric CO₂ concentrations (712 ppm) simulated together with climate change, did not prevent the decline in tree water use or soil water storage at the end of the summer. All scenarios caused the soil water storage to reach extremely low values at the end of the dry season (a minimum of 25 mm). It is concluded that climate change is likely to have a negative impact on tree water use and soil water resources in the study area, increasing the water deficit by as much as 30%.     

Regional mixed-effects height–diameter models for loblolly pine (<Emphasis Type="Italic">Pinus taeda</Emphasis> L.) plantations

A height–diameter mixed-effects model was developed for loblolly pine (Pinus taeda L.) plantations in the southeastern US. Data were obtained from a region-wide thinning study established by the Loblolly Pine Growth and Yield Research Cooperative at Virginia Tech. The height–diameter model was based on an allometric function, which was linearized to include both fixed- and random-effects parameters. A test of regional-specific fixed-effects parameters indicated that separate equations were needed to estimate total tree heights in the Piedmont and Coastal Plain physiographic regions. The effect of sample size on the ability to estimate random-effects parameters in a new plot was analyzed. For both regions, an increase in the number of sample trees decreased the bias when the equation was applied to independent data. This investigation showed that the use of a calibrated response using one sample tree per plot makes the inclusion of additional predictor variables (e.g., stand density) unnecessary. A numerical example demonstrates the methodology used to predict random effects parameters, and thus, to estimate plot specific height–diameter relationships.     

Effects of the interaction between drought and shade on water relations, gas exchange and morphological traits in cork oak (Quercus suber L.) seedlings

The combined effect of drought and light on different physiological and biochemical traits was assessed in cork oak (Quercus suber L.) seedlings grown under two levels of light availability and submitted to a long-standing drought. Watering was withdrawn after germination and seedlings were allowed to dry to a water content of ca. 50% of field capacity. At this point, water-stressed seedlings were grown under moderate drought and two light regimes: high light (HL—50%) and low light (LL—2%). Soil water in control plants was kept close to field capacity (90–100%) for both light environments. Water-relations parameters derived from P–V curves, gas exchange and water status at predawn (Ψ_pd) were evaluated at twice during the experiment. Nitrogen and chlorophyll contents were determined in the same leaves used for the gas exchange measurements. In addition, maximum rate of carboxylation (V_cmax) and electronic transport (J_max) were derived from A–C_i curves in well-watered seedlings.

The variation on moisture availability during the experiment was the same under both light environments. In control plants, Ψ_pd was over −0.3 MPa at the two harvests, while stressed seedlings decreased to −0.9 MPa, with no differences between light treatments. Water stress decreased osmotic potentials at full (Ψπ₁₀₀) and zero turgor (Ψπ₀). The regressions between both potentials and Ψ_pd showed a higher intercept in shade grown seedlings. This fact will point out the higher osmoregulation capacity in sun seedlings whatever water availability.

Nitrogen investment on a per leaf mass (N_mass), chlorophyll content (Chl_mass) and SLA tended to show a typical pattern of sun-shade acclimation. Thus, the three parameters increased with shade. Only for N_mass there was a significant effect of watering, since water stress increased N_mass.

LL plants showed a lower photosynthetic capacity in terms of maximum net photosynthesis at saturating light (A_max), which was related to a decrease in V_cmax and J_max. Both parameters varied with specific leaf area (SLA) in a similar way. The low-light environment brought about a higher nitrogen investment in chlorophyll, while under high-light environment the investment was higher in carboxylation (V_cmax) and electronic transport (F_max).

Stomatal conductance to water vapour (g_wv) and A_max were lower in low-light seedlings independently of watering. In addition, there was a trend to keep higher intrinsic water use efficiency (IWUE) under high light environment. The increase of IWUE under water stress was higher in HL seedlings. This was as consequence of the steeper decline in g_wv as Ψ_pd decreased. The decrease of A_max with Ψ_pd occurred in a similar way in LL and HL seedlings. Thus, the HL seedlings tended to sustain a higher ability to increase IWUE than LL seedlings when they were submitted to the same water stress.     

Effects of treeshelters and polyethylene mulch sheets on survival and growth of cork oak (Quercus suber L.) seedlings planted in northwestern Tunisia

Success of plantation establishment depends on the ability of seedlings to withstand animal browsing and competition from vegetation. This study in northwestern Tunisia examined the influence of browsing protection and vegetation control on seedling mortality, diameter, and height growth components of an evergreen sclerophyllous oak species, cork oak (Quercus suber L.) for the first 2 years after planting (2005 and 2006). Browsing protection was controlled by varying both treeshelter type (non-vented, vented, and control) and height (1.2-m and 1.8-m tall). Competing vegetation was controlled through increasing sizes of square-shaped black polyethylene mulch sheets: 0 m, 0.7 m, 1 m and 1.3 m large. Seedling mortality rate was relatively low during the first year (9%) and was not affected by any of the studied factors. During the following year, mortality was lower for seedlings in both types (non-vented and vented) of 1.2-m tall treeshelters than for unsheltered ones. Mortality was only slightly higher for 1.8-m tall shelters than for 1.2-m shelters. Compared to unsheltered seedlings, basal diameter was reduced during both seasons for seedlings inside non-vented treeshelters (1.2-m and 1.8-m tall), whereas for vented shelters it was reduced during the first year and then increased during the second year. Height growth was positively affected by tree shelters during both years. Seedlings inside non-vented 1.8-m tall shelters were the tallest having an average height of up to 140 cm at the end of the second year, while the unsheltered seedlings had only an average height of up to 42 cm. The greater height of the sheltered seedlings was attributed to the increase of: (1) the number of shoot flushes yearly established on the main stem (up to three), and (2) the length of all the growth units produced during each flush. Seedlings inside vented shelters had balanced growth with a low proportion of seedlings unable to support themselves without a stake (7%). However, non-vented shelters had unbalanced height and diameter growths resulting in both a higher height-to-diameter ratio and in a higher proportion of seedlings having stability problems (47%). Polyethylene mulch sheets did not affect early survival, seedling basal diameter, or number of shoot flushes established yearly, but did slightly improve seedling height at the end of the second season, mostly due to the significant improvement in shoot elongation of the first flush. Results suggest that both 1.8 m treeshelters and plastic mulches may enhance the growth of cork oak seedlings planted on harsh sites in northwestern Tunisia.     

A spatial distribution model of cork oak (Quercus suber) in southwestern Spain: A suitable tool for reforestation

Cork oak (Quercus suber) is an evergreen tree characterized by a thick bark, which grows in Mediterranean schlerophyllous forests. It is most prevalent in the southwestern Iberian Peninsula. Despite the potential of the province of Huelva (southwestern Spain) to maintain mature forests of cork oak, the tree has been severely depleted and most forests have either disappeared or are seriously threatened. This paper presents a spatial distribution model of cork oak for the province of Huelva with a view to determining the optimal areas for reforestation. The model draws on all available digital cartographic information with respect to cork oak distribution: topographic data (altitude, slope and orientation) were obtained from a Digital Terrain Model (20 m scale); rainfall, temperature and PET models were based on data collected from a network of meteorological stations; litologic data derive from the litologic map of Huelva (1:100,000). The result of this work is a mesh of points at a resolution of 100 m, sufficient to meet the needs of any kind of reforestation or management programmes in the area studied. Each point of this mesh contains the corresponding values for bioclimatic, topographic and litologic variables in a georeferenced data matrix. The independent variables responsible for cork oak distribution (binary dependent variables) were then identified by means of binary logistic regression analysis. North-facing slopes, abundant annual rainfall and litology were the main explaining variables. The spatial distribution model was produced by applying the formula obtained to spatial analysis software. This model is proposed as a basis for future reforestation plans, especially in those areas most affected by forest fires.     

Partial cutting as a conservation alternative for oak (Quercus spp.) forest—Response of bryophytes and lichens on dead wood

Semi-open oak-rich forests may support many species adapted to large trees and coarse woody debris. Currently many oak-rich forests in northern Europe have more or less closed canopies due to lack of natural and cultural disturbances. In these forests, conservation oriented partial harvesting of biofuel may restore a desired forest structure and light regime that potentially may favour many species that have declined during the last century. We quantified effects of such cutting on lichens and bryophytes living on dead wood in southern Sweden, using a design with paired cutting and control plots (each 1 ha) in 15 forests. In each plot we surveyed the same five logs and three stumps before and after harvest (25% of the tree basal area removed). The epixylic species composition shifted towards a flora typical for dryer dead wood. The mean species number of lichens per stump increased (clear effect) and the mean number of bryophytes per stump decreased (nearly significantly) due to harvesting. The species composition, but not species richness, on logs was affected. No increases or decreases in species of conservation concern could be detected. Since the overall negative effects were weak, we suggest that partial harvesting of dense oak forests is compatible with maintenance of the diversity of lichens and bryophytes on dead wood. However, the amount of such forest harvested needs to be carefully assessed in relation to demands of other taxa.     

PHENIPS—A comprehensive phenology model of Ips typographus (L.) (Col., Scolytinae) as a tool for hazard rating of bark beetle infestation

We developed the model PHENIPS for spatial and temporal simulation of the seasonal development of Ips typographus at the Kalkalpen National Park in Austria. The model is based on a digital elevation model used for interpolation of temperature and solar radiation to calculate the microclimatic conditions (bark temperature) for the beetles’ development. Additionally, the beetles’ phenology at Kalkalpen National Park was monitored along with air and bark temperature measurements. The onset of host tree infestation in spring was estimated using a lower threshold of 16.5 °C for flight activity and a mean thermal sum of 140 degree-days (dd) from beginning of April 1st onward. Rate of brood development was calculated from accumulated degree-days of hourly temperature data using upper and lower temperature thresholds of 38.9 and 8.3 °C, respectively, and a nonlinear function for calculating effective thermal sums. Re-emergence of parental beetles occurred at a time when 49.7% of the thermal sum for total development (557 dd) was reached. The model includes the discontinuance of the beetle's reproductive activity at a day length <14.5 h. The rate of successful hibernation of established broods is predicted by assessing the developmental stage of initiated generations at the beginning of the cold period. For validation we compared the timing of phenological events in the field with predicted events using both, hourly recorded data at trap trees in the terrain and generated daily topoclimatic data. Using topoclimatic data, the onset of infestation was predicted with a mean absolute error of 1.3 days. The observed onset of emergence of filial beetles in the field was estimated with a mean error of 39 dd. Our PHENIPS explicitly considers the strong effects of regional topography and stand conditions on local air and bark temperature and can be used for precise monitoring of the actual state of bark beetle development at the specific stand/tree level. Using topoclimatic data, PHENIPS simulates the maximum number of generations which is necessary to assess the potential impact of bark beetle outbreaks at regional scale. Further applications of PHENIPS for site-specific hazard rating of bark beetle infestation are discussed.