ࡱ>  wx]^|}k7 bjbjUU 7|7|Qd;#l$,RRRPS4NSD,%hVpWWW$WYN5ZQZ$ )aZY"YaZaZGqW$WߑGqGqGqaZ>RW$WGqaZGqtGqu:z,"{$WV )b,MRhfz {|0%zRpB{Gq,,GROWTH OF SIX CONIFEROUS SPECIES IN DIFFERENT BIOCLIMATES IN CROATIA SANJA PERI, STEVO ORLI, MLADEN IVANKOVI Forest Research Institute, Jastrebarsko, Cvjetno naselje 41, 10450 Jastrebarsko, Croatia, e-mail:  HYPERLINK "mailto:sanjap@sumins.hr" sanjap@sumins.hr Abstract Peri S., Orli S., Ivankovi M.: Growth of six coniferous species in different bioclimates in Croatia. Ekolgia (Bratislava), Vol. 23, Nr , , 2004. With the aim of determining a good selection of coniferous species for the establishment of cultures, the Forest Research Institute in Jastrebarsko began research work in 1960. Trial planting comprised three indigenous and three foreign species. The indigenous species included a Scotch pine (Pinus sylvestris L.), Black pine (Pinus nigra A r n.) and Norway spruce (Picea abies K a r s t.), and the foreign species were European Larch (Larix decidua M i l l.), Weymouth pine (Pinus strobus L.) and Douglas fir (Pseudotsuga menziesii var. viridis F r a n c o). These are economically the most interesting coniferous species that are often used in Croatia for establishing new cultures in non-forested areas and for reconstruction of lower quality stands of broadleaves. In spring of 1969 three comparative trials were established in three ecologically characteristic forest regions of Republic of Croatia. A necessary condition for successful production could simply be expressed in this way: "The right species in the right place". It can be achieved when the basic indicators of productive ability of stands as well as bio-ecologic traits of the species are known. An absolute success may be expected only in the cultures where the highest degree of harmonisation of species requirements regarding stands has been obtained. This paper shows the results of the 32-year development of plants of the aforementioned species. Key words: coniferous species, culture, survival, growth, height, DBH, wood volume, bio-ecological traits Introduction There are approximately 75 000 ha of coniferous cultures in Croatia. Another 330 000 ha of unoccupied forested and non-forested areas are available for possible forest harvesting. The largest areas available for establishing new coniferous cultures are located on the Mediterranean, Sub-Mediterranean and inland fern and heath areas (Mati et al., 1992). Continuous work on the selection of species and their provenance, as well as monitoring the success (growth and increment, as well as biomass production) of certain species in different edaphic, climatic and habitat conditions enables us to achieve better results in harvesting coniferous cultures. For successful production one needs to adhere to the principle  The right species in the right place (Orli, 1979). This can be implemented only if the biological i.e. ecological characteristics of the species as well as the basic indicators of the productive capability of the stand (soil quality) are known. During the last 40 years many scientists in Croatia have conducted research in existing cultures and have published many studies (Komlenovi et al., 1975; Komlenovi, 1976, 1978; Orli, 1979; Orli, Ocvirek, 1993; Dokua, Orli, 1986; Orli, Komlenovi, 1988). A large number of researchers from different countries also study this topic (Malknen, 1974; Lrgensen, 1976; Holmsgard, Bang 1977; Schrober, 1978; Hansen, Baker, 1979; Perie, Munson, 2000; Lindstrom, 2002; Maeunier et al., 2002). This paper shall demonstrate the results of research conducted on the success rate of six different species of conifers in comparative trials on three different locations in Croatia. Research Locations And Methods A series of comparative trials was set up in the spring of 1969 in three characteristic ecological forest areas in Croatia. The localities and their geographical position are given in Table 1 and Fig. 1. T a b l e 1. Geographical position of the localities Forest LocalityLatitudeLongitudeAltitude (m)Durgutovica Vinkovci 45 1918 38110Bjelovar, Slatki potok45461703142Karlovac, Lokve45261517195 The Lokve locality is situated in brake-tracts and heaths (Karlovac Forest Administration) in SW Croatia. The climate is perhumid (Bertovi, 1975). The Slatki potok locality is situated in NW Croatia in the hilly area of Bjelovar on agricultural land (Bjelovar Forest Administration). The climate is humid. The Durgutovica locality is situated in the East of Croatia on a once forested area (Vinkovci Forest Administration). The basic meteorological data is given in Table 2. T a b l e 2. Meteorological data per station Air temperature at 2 m [o C]Mean annualPrecipitation [mm] Station/MeanAmplitude of the AbsoluteAbsoluterel. airMean Min. annualMax. annualLocalityannualmean annual temp.max. temp.min. temp.humidity [%]annualsumsumKarlovac11.29.912.337.0-21.8771061.5896.51187.7Bjelovar10.89.811.836.9-18.880813.6655.2957.8Vinkovci11.810.514.236.7-21.082673.2470.0790.5  EMBED CorelDRAW.Graphic.10  Fig. 1. Position of the localities. The research encompasses the economically most interesting coniferous species (indigenous and foreign) that are most commonly used in Croatia for establishing new cultures outside forest areas and for reconstructing degraded broad-leaf stands. Planting material of known origin was used for the trial, with the same seed crop, identical nursery production and class of quality. The basic data on the origin and age of the seedlings is given in Table 3. T a b l e 3. Species and age of research seedlings SpeciesSymbolProvenanceAgeNorway spruce (Picea abeis K a r s t.)OSKlju , BiH2+2Scotch pine (Pinus sylvestris L.)OBViso , BiH1+2Black pine (Pinus nigra A r n.)CBTitovo u~ice,Yugoslavia1+2European Larch (Larix europea L.)EAKule, Brno, Czech R.1+1Weymouth pine (Pinus strobus L.)ABPonovec, Slovenia1+3Douglas-fir (Pseudotsuga menziesii F r a n c o)ZDVernonia Oregon, USA2+3 The trials were established in a randomised block design with three repetitions. Fig. 2 shows the layout of the trial. A total of 432 plants of every species (144 plants per lot) were planted during the trial, and the total number of plants per trial was 2591. The size of each trial including the protective cover equalled 1.5 ha. The planting distance was 2.0x2.0 m (2500 plant/ha). BlockIOSABEAZDOBCBIIZDOBCBOSABEAIIIABEAZDOBCBOS Fig. 2. Layout of the comparative trial. A series of research was conducted in these comparative trials: survival and height growth during the first five years, total height and breast height diameter by the end of the fifth year (Orli, 1979), height and width growth and increment and survival 10 years upon establishment (Orli, 1983), breast diameters and height after 23 and 26 years of age, stem wood, average and current increment, analysis of biomass and the concentration of nutrients in the plant matter (Komlenovi, 1978; Orli et al., 1991). This study gives the results of the latest measurement taken in the spring of 2002 for each of the three localities aged 32 years. The basic meteorological data is given for each locality. An analysis has been conducted regarding survival per species and locality. Total height and breast height diameter were measured on each tree. The wood volume and average age increment were calculated on the basis of the collected data. One entry tables tariffs were used for calculating wood volume. The statistical processing was conducted in Statistica and SAS 8.12 programmes. Descriptive statistic was done for all analysed variables. Difference between analysed species (Picea abies, Pinus sylvestris, Pinus nigra, Larix europaea, Pinus strobus, Pseudotsuga mensiesii) for height and DBH were testing using ANOVA for each location separately. If these difference was statistical significant ( p < 0.05) Scheffe test for multiple comparison were used for testing which species make this difference (Sokal, Rahlf, 1995). Research Results And Discussion Survival The survival rate of the plants was monitored in all the trials at 1, 10 and 32 years of age. The data is given in Table 4. T a b l e 4. Survival per species and locality SpeciesSum perDataYearOBCBOSEAZDABlocalityTrial: DurgutovicaPlanted (pc)19694324324324324324322592Survival [ %]196971889373808381Survival [ %]197858858469607070Survival [ %]200115357527496043Trial: Slatki potokPlanted (pc)19694324324324324324322592Survival [ %]1969869110096869692Survival [ %]197884858241167864Survival [%]200146537331165545Trial: LokvePlanted (pc)19694324324324324324322592Survival [ %]196981919990959892Survival [ %]197876749773659681Survival [%]20019167939345738Sum per speciesPlanted (pc)19691296129612961296129612967776Survival [ %]196979909786879288Survival [%]197873818861478172Survival [%]200123357632335742 As can be seen in the Table 4, the data on survival varies per year but nevertheless certain regularities can be established. Throughout the monitored years the highest survival rate is recorded in Norway spruce and Weymouth pine. A high survival rate is registered with the Black pine in the Durgutovica and Slatki potok localities (35%, 53%), while the low survival rate in the Lokve locality (16%) is the result of grave damage due to snowbreakage and wind breakage. The lowest survival rate throughout the trial period was recorded for the European Larch, Douglas fir and Scotch pine. If we are looking at survival rates per species for the year 2001 we can notice that it ranges from 9% for the Scotch pine on the Lokve locality to 79% for the Norway spruce at the same locality. The Weymouth pine and Norway spruce have a uniform survival rate on all three localities, which point to a wide ecological amplitude of these species. The European Larch and Douglas fir are species that havent shown tolerance in regard to locality selection and they have the lowest survival rate in 2001 ranging from 32% to 33%. A survival rate of only 23% for the Scotch pine is the result of values ranging from 9% to 46%. As regards locality, the locality with the highest survival rate of 45% was Slatki potok. Height growth Table 5 gives an overview of mean heights per tree species and locality. T a b l e 5. Mean heights and standard deviation Height [m]SpeciesDurgutovica Slatki potokLokveMeanStd. Dev. Mean Std. Dev.MeanStd. Dev.OB19.630.1719.530.1516.480.71CB17.810.0416.430.1114.600.67AB19.810.1322.140.3221.070.82ZD23.760.4121.120.8217.600.94EA21.020.3222.000.0321.020.49OS16.870.0217.720.2116.580.12 The European Larch (22.00 m) and Weymouth pine (22.14 m) are the leading species regarding height on the Slatki potok and Lokve localities. On the Durgutovica locality the Douglas-fir (23.76 m) is higher. The lowest heights were recorded for the Norway spruce and Black pine on all localities. This data corresponds to the data from earlier measurements (Orli, 1979, 1983). The relation between species is in accordance with their biological features and ecological requirements. The European Larch and Weymouth pine are superior species as regards the intensity of height growth during their young years in comparison to the Norway spruce and Black pine. Fig. 3 gives a graphical overview of the mean heights and a 95% reliability interval per species and locality. It is evident from the graph that the Black pine has had the smallest growth, and the narrowest range of heights was recorded with the European Larch. The data collected on height was analysed using ANOVA for each location separately. It has been statistically proven that significant differences exist in localities (Table 6). Scheffe test for multiple comparison were used for testing which species make this difference (Fig. 4). T a b l e 6. ANOVA for height and for each location separately SourceDFType I SS Mean Square F Value Pr > FSpecies-Lokve5103.588583320.717716744.71< 0.0001Species-Slatki potok583.4470277816.68940556116.84< 0.0001Species-Durgutovica589.3740944417.87481889331.66< 0.0001  EMBED STATISTICA.Graph \s  Fig. 3. Mean heights per species and locality. On the Lokve locality the species are divided according to the Scheffe multiple comparison test (Fig. 4) into three different groups. Significant differences exist between the Black pine and the other species. There is no major difference in height between the Waymouth pine and European Larch. These two species have the largest height values in this locality. The third group is composed of the Douglas fir, Norway spruce and Scotch pine. The graphical presentation shows the dominance of foreign species on this locality as regards height as well as the significant difference between the Douglas fir and the two other foreign species (Waymouth pine and European larch). Scheffe Grouping Mean N Species A 21.0733 3 3 L A o A 21.0167 3 5 k v B 17.6033 3 4 e B B 16.5800 3 6 B B 16.4767 3 1 C 14.6000 3 2 Scheffe Grouping Mean N Species S A 22.1400 3 3 l A a A 21.9967 3 5 t A k A 21.1233 3 4 i B 19.5267 3 1 p o C 17.7233 3 6 t o D 16.4267 3 2 k Scheffe Grouping Mean N Species D A 23.7633 3 4 u r B 21.0233 3 5 g u C 19.8067 3 3 t C o C 19.6267 3 1 v i D 17.8067 3 2 c a E 16.8700 3 6 Legend: Symbol of speciesSpecies1OB Scotch pine2CB Black pine3AB Weymouth pine4ZD Douglas fir5EA European Larch6OS Norway spruce Fig. 4. Scheffe multiple comparison test, heights On the other hand, on the Slatki potok locality there are no significant height differences according to the Scheffe test between trees of foreign species (Waymouth pine, Douglas fir, European larch). The indigenous species are significantly different from one another. Each indigenous species differs in height from all of the foreign species. On the Durgutovica locality the species are divided into five groups also in accordance with the Scheffe test. Only the Waymouth pine and Norway spruce dont have significant differences. All the other species have significantly differ in height. Diameter growth At age 32 the Douglas fir and Weymouth pine had the highest average breast height diameter. Data is given in Table 7 and Fig. 5. T a b l e 7. Average values for breast height diameter and standard deviation, cm Breast height diameter [cm]SpeciesDurgutovicaSlatki potokLokveMeanStd. Dev.MeanStd. Dev.MeanStd. Dev.OB22.370.7621.031.1422.032.40CB19.730.2118.600.5218.232.20AB20.530.7124.672.0522.972.74ZD23.131.4528.232.8720.502.50EA19.770.8522.100.1021.101.14OS16.370.0617.870.6817.372.40 Fig. 5. Mean DBH per species and locality The Douglas fir has the widest range within one species on all the monitored localities. On the Slatki potok locality all the species except for the Black pine and Norway spruce have shown the highest values of breast height diameters. It has been statistically proven that significant differences exist in breast height diameters depending on localities (Table 8). T a b l e 8. ANOVA for DBH and for each location separately SourceDFType I SS Mean Square F Value Pr > FSpecies-Lokve570.9333333314.186666673.310.0416Spec-Slatki potok5226.538333345.307666718.73< 0.0001Spec-Durgutovica585.2850000017.0570000025.91< 0.0001 The data was analysed using a single analysis variance (ANOVA) and tested with the Scheffe multiple comparison test (Fig 6). Scheffe Grouping Mean N Species A 22.967 3 3 L A o A 22.033 3 1 k A v A 21.100 3 5 e A A 20.500 3 4 A A 18.233 3 2 A A 17.367 3 6 Scheffe Grouping Mean N Species S A 28.233 3 4 l A a B A 24.667 3 3 t B k B C 22.100 3 5 i B C B C 21.033 3 1 p C o C 18.600 3 2 t C o C 17.867 3 6 k  Scheffe Grouping Mean N Species D A 23.1333 3 4 u A r B A 22.3667 3 1 g B A u B A C 20.5333 3 3 t B C o B C 19.7667 3 5 v C i C 19.7333 3 2 c a D 16.3667 3 6 Fig. 6. Scheffe multiple comparison test, DBH According to the Scheffe multiple comparison test for breast height diameters (Fig. 6) on the Lokve locality there are no significant differences between the six species subject to the research, which is also proven by the border value of F = 0.0416. On the Slatki potok locality the species were divided into three groups according to the same test. The foreign species Douglas fir and Waymouth pine had the largest values for breast height diameter that did not significantly differ from each other. The European larch and the Scotch pine do not significantly differ from the Waymouth pine, but they do differ from the Douglas fir, Black pine and Norway spruce. When comparing groups A, B and C that are the result of the Scheffe test, none of the indigenous species significantly differ from the European larch. The Scheffe test for breast height diameters on the Durgutovica locality shows four different groups. The Norway spruce significantly differs from the other species. It has the lowest values for breast height diameters on this locality. Apart from the Norway spruce, there are significant differences between the Scotch pine and Black pine, the Black pine and Douglas fir, and the Douglas fir and Norway spruce. Wood volume and basal area This study has also conducted an analysis of wood volume according to tree species and locality. The data is given in Table 9 and Fig. 7. T a b l e 9. Average values for wood volume and basal area SpeciesStem wood [m3/ha]Basal area [m2/ha]DSPLDSPLOB132.909376.15372.02815.366743.03719.4093CB267.874328.29981.87629.067538.516811.0114AB569.209731.028722.12954.233264.458963.9395ZD582.074250.522236.47551.787525.526427.1638EA229.492360.456414.49521.727634.110638.6164OS361.823422.172415.26540.433945.772346.7519  EMBED Excel.Chart.8 \s   EMBED Excel.Chart.8 \s  EMBED Excel.Chart.8 \s  Fig. 7. Overview of wood volume per tree species and locality The overview illustrates that the species with the most wood volume in all three localities is the Weymouth pine. The wood volume ranges from 569 m3/ha to 731 m3/ha. On the Durgutovica locality along side the Weymouth pine, the Douglas fir has the highest wood volume value. The Scotch pine (72 m3/ha) and Black pine (81 m3/ha) have an extremely low wood volume value on the Lokve locality. These results are related to the earlier mentioned snow breakage and wind breakage on this locality. On the Slatki potok locality, most of the monitored species (every species except the Douglas-fir) have had the highest wood volume values. When taking into consideration all the factors i.e. survival rate, height, breast height diameter and wood volume we can generally conclude that all the species grow best on the Slatki potok locality. Conclusions On the basis of the research conducted on the six different conifer species in their 32nd year over three localities we can conclude the following: Throughout the monitored years (1,5,10,32) the highest survival rate was recorded for Norway spruce and Weymouth pine, and the lowest for European Larch, Douglas fir and Scotch pine. The Weymouth pine and Norway spruce have a harmonized survival rate on all three localities. This indicates wide ecological amplitude of these species. The European larch and Weymouth pine have the highest height values on the Slatki potok and Lokve localities, and on the Durgutovica locality they follow the values for the Douglas fir. The lowest values for height pertain to the Norway spruce and Black pine on all localities. Statistical analysis has proven significant differences in breast height diameters and heights depending on tree species and locality, as well as the existence of interaction between locality and species. The highest values for breast height diameter were in average achieved by Douglas fir and Weymouth pine, and the lowest by Norway spruce and Black pine. The Weymouth pine has the highest average wood volume value on all three localities. The Slatki potok locality has proven to be the best as regards growth success for all six species of conifers. Different climatic conditions and productive capabilities of the stands as well as biological characteristics influence individual species on different localities. 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