A+ A A-

Short-term effect of early thinning on growth in stone pine in Tunisia

Download this file (JNS_AgriBiotech_Vol_46_02.pdf)Volume 46, Article 02[Volume 46, Article 02]296 kB

Effet à court terme d’une éclaircie précoce sur la croissance de Pin pignon en Tunisie





Sciences faculty of Bizerte (FSB), Jarzouna - 7021- university of carthage -Tunisia.

2 Silvopastoral Institute —Tabarka, laboratory of silvopastoral resources. University of Jendouba.

3 Department of Sylviculture and Management of Forest Systems (INIA-CIFOR). Ctra. de La Coruňa, Km. 7,5. 28040 Madrid- Spain.


Abstract – The aim of this study was to quantify growth responses of 14-year-old artificially regenerated stone pine (Pinus pinea L.) stands to thinning in Northwestern Tunisia, four years after treatment. The study included four thinning intensities (light (17% of trees was removed), moderate (40% of trees was removed), strong (63% of trees was removed) and very strong (81% of trees was removed) thinning), plus control (0% of trees was removed) replicated in four randomized block design. The variables measured included height growth, diameter at breast height (dbh), crown radius, branch diameter of the whorl closest to breast height (1.3 m), length and dry mass of needles, and soil water content. Thinning led to an increase in dbh between 3 and 35%, while there was no significant effect on height growth. Thinning also promoted crown expansion and branch diameter, whereas no effects on length and dry mass of needles were found. On the other hand, there were no differences among treated and untreated plots with regard to soil water content. Thus, changes in tree growth tend to be linked most closely to increases in space and light availability in the thinned plots.

 Keywords: thinning; Pinus pinea L.; growth

Résumé - L’objectif de la présente étude est de quantifier la réponse de croissance d’un peuplement jeune (14 ans) de Pin pignon (Pinus pinea L.), régénéré artificiellement, à l’éclaircie. Quatre intensités d’éclaircie (faible (en enlevant 17% des arbres), modérée (en enlevant 40% des arbres), forte (en enlevant 63% des arbres) et très forte (en enlevant 81% des arbres)) en plus d’un témoin (non touché), répétés en quatre blocs randomisés, ont été ainsi appliqués. L’étude a été menée, durant 4 ans 
(1994-1998), au Nord-Ouest de la Tunisie. Les mesures réalisées ont porté sur la croissance en hauteur, le diamètre à la hauteur de poitrine (1,3 m), le rayon du houppier, le diamètre des branches du verticille le plus proche de 1,3 m, la longueur et la biomasse sèche des aiguilles ainsi que le bilan hydrique. Quatre ans après l’éclaircie, le diamètre des arbres a augmenté de 3 à 35%, alors que le changement en hauteur a été insignifiant. L’effet de l’éclaircie s’est traduit également par une augmentation de la largeur du houppier et du diamètre des branches. En revanche, aucun effet significatif sur la croissance en longueur ou en biomasse des aiguilles n’a été trouvé. L’éclaircie n’a pas eu d’effet significatif aussi sur le bilan hydrique, et les changements de croissance notés semblent être donc étroitement liés à l’augmentation de l’espace et de la disponibilité de la lumière autour des arbres des parcelles traitées.

Mots clés : éclaircie ; Pinus pinea L. ; croissance

1. Introduction 

Stone pine (Pinus pinea L.) is a species native to the Mediterranean area, where it covers more than 700,000 ha (Mutke et al. 2012), mainly in Spain (450,000 ha), Portugal (90,000 ha), Turkey (50,000 ha) and Italy (40,000 ha) (Pereira et al. 2015). It was successfully introduced in Tunisia more than a century ago (1907) being used primarily, along the Mediterranean coast line, to consolidate the coastal dunes of Bizerte in the North and along the North East coast in the region of Cap Bon (Hasnaoui 2000). The success of these first plantations incited the foresters to use this species to stabilize the coastal dunes of the Northwest too. Today, Pinus pinea occupies an area of 21,165 ha

(El Khorchani 2010) and becomes one of the most valuable species in Tunisian reforestation programs due to its usefulness for both wood and fruit production (pine nuts) and its ability to grow in dry and sandy soils. These stone pine forests suffer, however, a lack of silvicultural interventions that can potentially aid in improving their growing conditions.

Thinning is the most commonly applied silvicultural treatment, thus it has received considerable attention in forest research. Thinning has traditionally been used to increase tree growth or improve its quality on a sustainable basis (e.g. Houtzagers 2002; Tullus 2002; Zeide 2004). After thinning, remaining trees can use the released resources, such as light, water and nutrients, to expand their crowns and to grow faster in diameter (Mäkinen and Isomäki 2004a; González-Ochoa et al. 2004; Crecente-Campo et al. 2009; Pérez-de-Lis et al. 2011). When properly used, thinning reduces

long-term stress by competition, but it also reduces the vulnerability of trees to extreme drought events (Linares et al. 2011; Sánchez-Salguero et al. 2012). However, the advantages of thinning for growth may not be the same for trees of different species (Moreno-Fernández et al. 2014; Navarro-Cerrillo et al. 2016) depending on their shade tolerance (Navarro-Cerrillo et al. 2016).

In the Mediterranean basin, several thinning experiments have been carried out in other pines (e.g. Del Río et al. 2008; Montero et al. 2001; Navarro-Cerrillo et al. 2016). However, there is much less information about the effect of thinning on stone pine (Gordo et al. 2009), which contrasts with the utility of the species. Thus, the main objective of this study was to analyze the effects of thinning on the growth of a 14-years-old artificially regenerated stone pine stand, four years after treatment application.

2.Matériel et méthodes

2.1. Study area and vegetal material

The study was carried out between 1994 and 1998, on a14-years-old low pole stone pine stand, at the plot 8 of the first serial of Mekna I (36°57′3.88′′N, 8°48′49.98′′E, 48 m a.s.l) in Northwestern Tunisia. The plot area has5 ha, and was occupied by eucalyptus until 1980, when it was substitute with a stone pine plantation. The climate is Mediterranean with an annual mean temperature of 17.9°C (1900 - 1988) (Hasnaoui 1992). Absolute maximum and minimum temperatures are 47°C (August) and -1°C (January), respectively (Hasnaoui 1992). The average annual rainfall is 1013 mm, with 46% of the total rainfall falling in winter, 24% in spring, 3% in summer and 27% in autumn.

2.2. Experimental design and treatments

The experiment was arranged as a randomized complete block with four replications 
(or blocks). The thinning treatments, in each block, were carried out on five experimental plots: control plot (0% of trees was removed); plot with light thinning where 17% of trees was removed; plot with moderate thinning where 40% of trees was removed; plot with strong thinning where 63% of trees was removed, and plot with very strong thinning where 81% of trees was removed. Each treatment was applied in one 400 m
2 plot. Thinning treatments were carried out in 1994, and removed trees were trees with smaller diameters and tree forks.

2.3. Measurements

Tree height, diameter at breast height (dbh) and crown radius were measured just after thinning (1994). They were recorded again at the end of the experiment (1998) along with tree density (number of remaining trees per treatment after thinning), branch diameter and soil water content. Measurements of diameter were systematically made, by keeping the same direction of measurement as well as the same height (1.30 m). Ten trees of a mean arithmetical diameter were selected from each treatment for height measurements. Measurements were made using a ranging rod. When height exceeds 6 meters we use the Blume Leiss. On the 10 trees previously selected, we measured three radii of the crown by projection on the soil. These radii were spaced of 120° approximately, and the average radius was calculated. Branch diameter was measured in the first whorl closest to the breast height

(1.3 m). The average branch diameter was obtained by dividing the sum of the branch diameters by the number of branches. On the same selected trees, 30 needles (3 needles per tree) aged two years old (1996-1998) were removed and measured for their length. Fresh and dry needle (at 60°C for 48 hours) weight was calculated. To appreciate the water conditions of the experimental plot, four samples of soil per treatment were taken, in spring 1998, at 40 cm depth. They were then mixed to make a composite sample in each plot (or treatment). This volume was weighed in a fresh state then in a dry state after drying at 100°C for 48 hours to determine the soil moisture rate.

2.4. Data analysis

Data were analyzed using the Statistical Analysis System (SAS Institute Inc., Cary, NC), corresponding to the following general linear model (Anon 1998):

Yij = µ + Ai + Bj + eij

Where Yij = the independent variable; µ = general mean; Ai = effects of block; Bj = effects of thinning treatments and eij = error term.

For each analysis, when the analysis of variance was significant, statistically significant differences between means were identified using Waller-Duncan K-ratio = 100 (Little and Hill 1978). Differences were considered significant at P ≤ 0.05. Finally, all measured parameters were used to make an analysis of the correlation by using the test of Pearson correlation.

3. Results and discussion

3.1. Tree characteristics at establishment of the experiment

At the beginning of the experiment, just after thinning (1994), height, diameter at breast height (dbh), and crown radius of trees were similar between thinning treatments (Table 1). By contrast, significant differences in tree density between thinning treatments were found. Hence, any future change in trees growth between thinning treatments from then on would be interpreted as it is due to the thinning operation.


Table 1. Analysis of variance (Pr>F) and mean values of tree density, diameter at breast height (1.3 m), tree height and crown radius, just after thinning application (1994).







Very strong









Density (number of trees/treatment)







Diameter at 1.3 m (cm)







Tree height (m)







Crown radius (m)








For a same line, P values marked with an asterisk (*) indicate the presence of a significant thinning effect, while mean values marked with different letters indicate the presence of a significant difference at P ≤ 0.05 level.


3.2. Radial growth
Studies on the effect of thinning in diameter growth of residual trees are numerous for many tree species from a wide geographical range. These studies demonstrated that thinning promotes dbh of many species of both conifer and hardwood trees (Briggs and Lemin 1994; Burns et al. 1996; Varmola et al. 2004; Medhurst et al. 2001; Juodvalkis et al. 2005; Rytter and Werner 2007). This is in agreement with the results of our work which show that thinning favored diameter growth; the trees which initially (just after thinning, 1994) had no significant differences in diameter at breast height (dbh) showed after four years of growth (1998) statistically significant differences between thinning treatments, and average dbh was greater for thinned than for unthinned (control) plots by 3% (0.4 cm), 10% (1.3 cm), 17% (2.2 cm), 35% (4.4 cm), respectively under light, moderate, strong and very strong thinning (Table 2). Differences were, however, significant only for strong and very strong intensities of thinning indicating that thinning effect is influenced by thinning intensity. On the other hand, while both of these thinning regimes resulted in a significantly increase in diameter growth this diameter was significantly greater under very strong than under strong thinning. This is in line with other results obtained in many studies carried out in central and northern Europe which establish that thinning intensity must be high (e.g. Mäkinen and Isomäki 2004b). It has been shown recently, however, that responses of tree species to thinning depend on their shade tolerance; for instance, Pinus sylvestris showed a significantly higher radial growth under light thinning intensity, while Pinus pinaster which it is less tolerant to shade showed a significantly higher radial growth under heavy thinning

(Navarro-Cerrillo et al. 2016). P. pinea is reputed to be a very light-demanding species (Adili 2012), which could explain why radial growth reached its maximum under very strong thinning regime.

3.3. Tree height and needles growth

Contrary to trees diameter, height was unaffected by thinning intensity (Table 2). Similar results were found in various broad-leaved (Graham 1998; Medhurst et al. 2001; Rytter and Werner 2007; Çiçeket al. 2013; Diaconu et al. 2015) or coniferous (Del Río et al. 2008; Mäkinen and Isomäki 2004b; Crecente-Campo et al. 2009) species. It was reported that stand density has significant effects on diameter growth but not on height growth, except for very high and very low stand densities (Çiçeket al. 2013). In our study thinning did not affect height growth, meaning that tree densities are at a level that does not affect trees height. On the other hand, it has been also reported since long that the carbon allocation for height growth is more primary than the carbon allocation for diameter growth (Lanner 1985) which may also explain why thinning did not affect height growth. As shown in

table 2, there were also no significant differences between thinned and unthinned plots neither for length nor for the dry mass of needles. It is known, in general, that morphological needle parameters increase with increasing tree height (Gebaueret al. 2011) reason for which perhaps both length and dry mass of needles, as found for height growth, was independent of thinning treatments.

3.4. Branch diameter

It is well known that tree branches result in knots within the stem, which reduces strength because of changes in fibre angles and interruption of wood continuity as growth rings of branches join growth rings of the stems (Raprager 1939; Green et al. 1999). In addition, compression wood is formed at the base of branches via adaptive responses that increase support for them (Schultz 1997). Both of these traits negatively influence wood quality, especially on strength and shrinkage parameters (Perstorper et al. 2001). Similar to other studies (e.g. Varmola and Salminen 2004; Fahlvik et al. 2005), our study showed that the mean branch diameter of the whorls closest to breast height tended to increase with increasing thinning intensity (Table 2). The result is, therefore, increased diameter knots within the stem. Accordingly, retention of branches on the stem after thinning, and especially under very strong thinning regime where branch diameter was found to be significantly increased compared to unthinned control plots as our results show, is not recommended when thinning is oriented towards the production of high-quality timber.

3.5. Crown projection and soil water content

Juodvalkis et al. (2005) showed that a very significant rise in the increase of crown volume can be achieved through thinning in the case of young trees – for example, those aged 10–20 years in the case of pines. This may be confirmed in our study for stone pine, where thinning was found to increase crown radius, on average, by 11 to 28% although the difference under light thinning was not significant compared to unthinned control plots (Table 2). Crown width expansion in remaining trees suggests that trees tended to maximize their space and light interception. It was found, on the other hand, for the same species that tree-level cone production is positively related to diameter and crown width (Calama et al. 2008; Gonçalves and Pommerening 2012). So when the management goal is fruit production, thinning can be done to increase crown growth which will also increase fruit production. Based on the results obtained for crown and diameter growths we can establish that trees had a higher dbh increase (there was a significant increase in dbh under very strong thinning regime compared to strong thinning regime) rather than crown width increase (there were no significant differences between moderate, strong and very strong thinning regimes (see table 2)). This is because trees, as reported by Loewe et al. (2013) seem to devote their resources to consolidate first, and later to a greater fruit production. It is interestingly to emphasize, on the other hand, that increased crown growth under moderate thinning and its stabilization since, suggest that this thinning regime is largely sufficient to stimulate crown development, when the forest objective is the fruit production, which may help to minimize the cost of the thinning operation comparatively to strong and very strong thinning regimes.

Several thinning studies have reported that in Mediterranean sites, thinning could enhance tree growth by improving the tree water status (Bréda et al. 1995; Ma et al. 2010; Molina and Del Campo 2012), since tree growth is usually mainly limited by water deficit (Sabaté et al. 2002). However, more recently Primicia et al. (2016) found that soil moisture was higher in unthinned than in thinned plots even 9 years after the first thinning. In our study plots we did not observe also, after four years, that thinning increases soil moisture and differences between treatments were not significant (Table 2). These contrasting results indicate that thinning does not guarantee the improvement of water availability in thinned plots. Finally, it should be underlined that the absence of a soil water content effect of thinning found in this study could indicate that in thinned plots ilow tree density make better use of available water (by expanding roots) and ii) there was an increase of evapotranspiration.


Table 2. Analysis of variance (Pr>F) and mean values of diameter at breast height (dbh), tree height, branch diameter, crown radius, length and dry mass of needles and soil water content under different thinning intensities: control, light thinning intensity, moderate thinning intensity, strong thinning intensity and very strong thinning intensity.






Very strong

Pr> F








dbh (cm)







Tree height (m)







Branch diameter (cm)







Crown radius (m)







Needles length (cm)







Dry mass of needles (g)







Soil water content (%)







For a same line, P values marked with an asterisk (*) indicate the presence of a significant thinning effect, while mean values marked with different letters indicate the presence of a significant difference at P ≤ 0.05 level.

3.6. Correlation between variables
The correlation analysis, presented in table 3, clearly demonstrates that the growth traits significantly affected by thinning (dbh, crown projection, branch diameter) were significantly correlated with tree density, which was not the case for those growth traits that were independent of thinning effect (height growth, length and mass of needles). These results support the general idea that tree growth is regulated by thinning by controlling stand density (Mäkinen and Isomäki 2004a). For the same species (Pinus pinea L.), diameter growth was found to be positively correlated with crown size (Ciancio et al. 1986). However, in our study we did not find that crown expansion is correlated with diameter growth; hence, the larger trees, the larger crown seems to be not the case in this study, and crown width seems to be more proportional to tree height than to tree diameter, as it significantly and positively correlated with height growth. Irrespective of its correlation with tree density, branch diameter was found to be significantly and positively correlated with dbh which is in agreement with the results documented by other researchers for other coniferous species (e.g. Persson 1977 for Scot pine; Pfister 2007 for Norway spruce; Liziniewicz 2014 for Norway spruce, Scots pine, Lodgepole pine). Finally, it should be underlined that the dependence of needle morphology on height growth, as previously mentioned, may be confirmed but only for needle length which was found to be significantly correlated with tree height; needle mass was not correlated with tree height, but rather with needle length.


Table 3. Pearson’s correlation (R2) between different variables, with T.D = tree density, dbh = diameter at breast height 
(1.3 m), H = tree height, B.D = branch diameter, C.R = crown radius, N.L = needle length, N.M = needles mass.









T.D (trees/plots)

























































Values marked with an asterisk (*) indicate the presence of a significant correlation at P ≤ 0.05 level.

4. Conclusion
The results from our study show that thinning may have a positive effect on growth of young stone regenerated artificially pine stands. This effect was, however, dependent on thinning intensity. Thus, trees started to react significantly under moderate thinning regime for crown expansion, under strong thinning regime for dbh, and under very strong thinning regime for branch diameter. These observed growth changes under thinning seemed to be not linked to soil moisture, which was not improved after thinning, but rather to increase in the available space surrounding trees and the amount of intercepted light. It is known that tree branches become knots within the stem. Thus, retention of branches on the stem under very strong thinning, where there was a significant increase in diameter of these branches, may result in increases in both the abundance and size of knots. However, knots within the stem constitute a defect when the tree is assessed for timber quality and when the tree is processed into boards. In this case, early prunings under very strong thinning are therefore required. Contrary to diameter growth, thinning did not lead to an increase in height growth. Thus, when we consider the height growth as an index of site quality one can draw the conclusion that there were no differences in the fertility of different plots, and that thinning did not create therefore fertility classes. In general, our results allows to gain important knowledge about the effects of different thinning regimes in stone pine forests, which will contribute to improve stone pine forest management.


This research supported by the laboratory of silvopastoral resources (Silvopastoral Institute — Tabarka). The authors would like to thank the Tunisian General Direction of Forests (DGF) and their staff in the district of Tabarka for permission to conduct the study and for assistance in the field. Special thanks to Dr. Ali AlOUI for providing technical assistance.

5. References

Adili B (2012) Croissance, fructification et régénération naturelle des peuplements artificiels de Pin pignon (Pinus pinea L.) au nord de la Tunisie. Thèse de doctorat, Université Blaise Pascal - Clermont-Ferrand II.

Anon (1998) SAS/STATTM User’s Guide. Release 6.03 edn. SAS Institute, Cary, NC.

Breda N, Granier A, Aussenac G (1995) Effects of thinning on soil and tree water relations, transpiration and growth in an oak forest (Quercus petraea Matt.) Liebl.). Tree Physiol 15: 295–306.

Briggs RD, Lemin RC (1994) Soil drainage class effects on early response of balsam fir to 
pre-commercial thinning. Soil Sci Soc Am J 58(4): 1231-1239.

Bums J, Puettmann KJ, Perala D (1996) Strip thinning and spacing increases tree growth of young black spruce. North J Appl For 13(2): 68-72.

Calama R, Mutke S, Gordo J, Montero G (2008) An empirical ecological type model for predicting stone pine (Pinus pinea L.) cone production in the Northern Plateau (Spain). For Ecol Manage 255:660–673. doi:10.1016/j.foreco.2007.09.079.

Ciancio O, Cutini A, Mercurio R, Veracini A (1986) Sulla struttura della pineta di pino domestico di Alberese. Ann Ist Sper Selv Arezzo 17:169–236.

Çiçek E (2013) Effects of thinning intensity on the growth of narrow-leaved ash (Fraxinus angustifolia subsp. oxycarpa) plantations. Turk J Agric For 37: 97-104. doi:10.3906/tar-1204-13.

Crecente-Campo F, Pommerening A, Rodríguez-Soalleiro R (2009) Impacts of thinning on structure, growth and risk of crown fire in a Pinus sylvestris L. plantation in northern Spain. For Ecol Manage 257: 1945–1954.

Del Río M, Calama R, Cañellas I, Roig S, Montero G (2008) Thinning intensity and growth response in SW-European Scots pine stands. Ann For Sci 65:308p1-308p10. doi: 10.1051/forest: 2008009.

Diaconu D, Kahle HP, Spiecker H (2015) Tree- and Stand-Level Thinning Effects on Growth of European Beech (Fagus sylvatica L.) on a Northeast- and a Southwest-Facing Slope in Southwest Germany. Forests 6: 3256-3277. doi:10.3390/f6093256.

El Khorchani A (2010) The Pinus pinea. Forests in Tunisia. AGORA, International Scientific Workshop for Young Researchers, Tunisia (Hammamet).

Fahlvik N, Ekö P-M, Pettersson N (2005) Influence of precommercial thinning grade on branch diameter and crown ratio in Pinus sylvestris in southern Sweden. Scand J For Res 20: 
243 – 251.

Gonçalves AC, Pommerening A (2012) Spatial dynamics of cone production in Mediterranean climates: a case study of Pinus pinea L. in Portugal. For Eco Manag 266:83–93. doi:10.1016/j.foreco.2011.11.007.

González-Ochoa AI, López-Serrano FR, de las Heras J (2004) Does postfire forest management increase tree growth and cone production in Pinus halepensisFor EcolManage 188:

Gordo J, Calama R, Rojo LI, Madrigal G, Álvarez D, Mutke S, Montero G, Finat L (2009) Experiencias de clareos en masas de Pinus pinea L. en la Meseta Norte. In: Summaries of the Spanish fifth forestry congress. S.E.C.F.-Junta de Castilla y León.

Graham JS (1998) Thinning increases diameter growth of paper birch in the Susitna Valley, Alaska: 20 year results. North J Appl For 15: 113–115.

Green DW, Winandy JE, Kretschmann DE (1999) Mechanical Properties of Wood. In: Wood handbook: wood as an engineering material. Gen. Tech. Rep. FPL–GTR–113. Madison, WI: USDA Forest Service, Forest Products Laboratory. Chapter 4: pp 1–44.

Hasnaoui F (2000) Sciage et séchage du Pin pignon: propriétés physiques et mécaniques. Mémoire de PFE, INAT, Tunisie.

Houtzagers MR (2002) Thinning in discussion (in Dutch). Nederlands Bosbouwtijdschr, 74:7–14.

Juodvalkis A, Kairiukstis L, Vasiliauskas R (2005) Effects of thinning on growth of six tree species in north-temperate forests of Lithuania. Eur J Forest Res 124: 187–192. doi 10.1007/s10342-005-0070-x.

Lanner RL (1985) On the sensitivity of height growth to spacing. For Ecol Manage 13: 143–148.

Linares JC, Delgado-Huertas A, Carreira JA (2011) Climatic trends and different drought adaptive capacity and vulnerability in a mixed Abies pinsapoPinus halepensis forestClimatic Change: 105: 67–90.

Little TM, Hills FJ (1978) Agricultural Experimentation: Design and Analysis. John Wiley and Sons, New York, NY.

Liziniewicz M (2014) Influence of spacing and thinning on wood properties in conifer plantations. Doctoral Thesis Swedish University of Agricultural Sciences.

Loewe V, Venegas A, Delard C, González M (2013) Thinning effect in two young stone pine plantations (Pinus pinea L.) in central southern Chile. In: Mutke S (ed) Piqué M (ed), Calama R (ed) Mediterranean stone pine for agroforestry. Zaragoza : CIHEAM / FAO / INIA / IRTA / CESEFOR / CTFC, pp 49 -55 (Options Méditerranéennes : Série A. Séminaires Méditerranéens; n. 105).

Ma SY, Concilio A, Oakley B, North M, Chen JQ (2010) Spatial variability in microclimate in a mixed-conifer forest before and after thinning and burning treatments. For Ecol Manage259: 904–915. http://dx.doi.org/10.1016/j.foreco.2009.11.030.

Mäkinen H, Isomäki A (2004a) Thinning intensity and growth of Norway spruce stands in Finland. Forestry (Oxford) 77:349–364.doi:10.1093/forestry/77.4.349.

Mäkinen H, Isomäki A (2004b) Thinning intensity and growth of Scots pine stands in Finland. For Ecol Manage 201:311–325.doi:10.1016/j.foreco.2004.07.016.

Medhurst JL, Beadle CL, Nielson WA (2001) Early-age and later-age thinning affects growth, dominance, and intraspecific competition in Eucalyptus nitens plantations. Can J Forest Res 31: 187–197.

Molina AJ, Del Campo AD (2012) The effects of experimental thinning on throughfall and stemflow: A contribution towards hydrology-oriented silviculture in Aleppo pine plantations. For Ecol Manage 269: 206–213.

Montero G, Calama R, Ruiz-Peinado R (2008) Selvicultura de Pinus pinea L. In: Serrada R, Montero G, Reque JA (eds). Compendio de Selvicultura Aplicada en España. INIA-Ministerio de Educación y Ciencia. Madrid, Spain 1178 pp.

Moreno-Fernández D, Sánchez M, Álvarez JG, Hevia A, Majada JP, Cañellas I, Gea-Izquierdo G (2014) Response to the interaction of thinning and pruning of pine species in Mediterranean mountains. Eur J For Res 133: 833–843.

Mutke S, Calama R, González-Martinez S, Montero G, Gordo J, Bono D, Gil L (2012) Mediterranean stone pine: botany and horticulture. Horticultural Reviews 39: 153–202.

Navarro-Cerrillo RM, Sánchez-Salguero R, Herrera R, Ceacero Ruiz CJ, Moreno-Rojas JM, Manzanedo RD, López-Quintanilla J (2016) Contrasting growth and water use efficiency after thinning in mixed Abies pinsapo – Pinus pinaster – Pinus sylvestrisforests. J For Sci 62(2): 53–64. doi: 10.17221/104/2015-JFS.

Pereira S, Prieto A, Calama R, Diaz-Balteiro L (2015) Optimal management in Pinus pinea L. stands combining silvicultural schedules for timber and cone production. Silva Fennica 49 (3): 16 p.

Pérez-de-Lis G, García-González I, Rozas V, Arévalo JR (2011) Effects of thinning intensity on radial growth patterns and temperature sensitivity in Pinus canariensis afforestations on Tenerife Island, Spain. Ann For Sci 68 1093–1104.

Persson A (1977) Quality development in young spacing trials with Scots pine. Stockholm: Department of forest yield research, Royal College of Forestry. Research notes 45. Report no 45.

Perstorper M, Johansson M, Kliger R, Johansson G (2001) Distortion of Norway spruce timber - Part 1. Variation of relevant wood properties. Holz Als Roh-Und Werkstoff 59(1-2): 94-103.

Pfister O, Wallentin C, Nilsson U, Ekö PM (2007) Effects of wide spacing and thinning strategies on wood quality in Norway spruce (Picea abies) stands in southern Sweden. Scand J For Res 22(4): 333-343.

Primicia I, Artázcoz R, Imbert JB, Puertas F, María-del-Carmen Traver MDC, Federico-José Castillo FJ (2016) Influence of thinning intensity and canopy type on Scots pine stand and growth dynamics in a mixed managed forest. Forest Systems, 25(2), e057.

Raprager EF (1939) Development of branches and knots in western white pine. Journal of Forestry3: 239-245.

Rytter L, Werner M (2007) Influence of early thinning in broadleaved stands on development of remaining stems. Scand J For Res 22: 198–210.

Sabaté S, Gracia CA, Sánchez A (2002) Likely effects of climate change on growth of Quercus ilexPinus halepensisPinus pinasterPinus sylvestris and Fagus sylvatica forests in the Mediterranean region. For Ecol Manage 162: 23–37.

Sánchez-Salguero R, Navarro-Cerrillo RM, Swetnam TW, Zavala MA (2012) Is drought the main decline factor at the rear edge of Europe? The case of southern Iberian pine plantations. For Ecol Manage 271: 158–169.

Tullus H (2002) The influence of intermediate cuttings on the growth of pine and spruce forests: silvicultural recommendations. Metsanduslikud Uurimused (Forestry Studies, Tartu) 36:

Varmola M, Salminen H (2004) Timing and intensity of precommercial thinning in Pinus sylvestris stands. Scand J For Res 19: 142 – 151.

Varmola M, Salminen H, Timonen M (2004) Thinning response and growth trends of seeded Scots pine stands at the arctic timberline. Silva Fennica 38(1): 71–83.

Zeide B (2004) Optimal stand density: a solution. Can J For Res 34:846–854.



This article is published under license to Journal of New Sciences. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

CC BY 4.0