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Foxtailing of Pinus caribaea var. hondurensis in peninsular Malaysia : frequency, growth rate and specific… Ibrahim, Zakaria 1984

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FOXTAILING OF PINUS CARIBAEA VAR. HONDURENSIS IN PENINSULAR MALAYSIA: FREQUENCY, GROWTH RATE AND SPECIFIC GRAVITY by (5) ZAKARIA IBRAHIM B.Sc. (For) Universiti Pertanian Malaysia A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Forestry in The Faculty of Graduate Studies (Faculty of Forestry, The University of British Columbia) We accept this thesis as conforming to the required standard The University of British Columbia January 1984 © Zakaria Ibrahim, 1984 In presenting t h i s thesis i n p a r t i a l f u l f i l m e n t of the requirements f o r an advanced degree at the University of B r i t i s h Columbia, I agree that the Library s h a l l make i t f r e e l y a v a i l a b l e for reference and study. I further agree that permission for extensive copying of t h i s thesis f o r s c h o l a r l y purposes may be granted by the head of my department or by h i s or her representatives. I t i s understood that copying or publ i c a t i o n of t h i s thesis f o r f i n a n c i a l gain s h a l l not be allowed without my written permission. (ZAKARIA IBRAHIM) Department of F o r e s t r y The University of B r i t i s h Columbia 1956 Main Mall Vancouver, Canada V6T 1Y3 Date A p r i l 1 1 , 1984. DE-6 (3/81) i ABSTRACT Foxtailing is a common feature in the plantations of Pinus  caribaea var. hondurensis Barrett and Golfari in Peninsular Malaysia. Frequency of foxtailing in Kemasul and Ulu Sedeli pine plantations, aged between 1 to 8 years, was found to vary between 4.3 to 36.0 per-cent. Ulu Sedeli plantation has 5.3 percent more foxtail than in Kemasul plantation. This study indicates that the occurrence of fox-tailing varies with site and age. The most common form of foxtailing is the sub-terminal foxtail which constitutes about 60.0 percent of the foxtail population. The increasing proportion of sub-terminal to terminal foxtail with age of the trees suggests that foxtailing is a plastic trait. Average total height of foxtailed trees was greater than normal trees at all ages, however, larger diameters were evident only during the juvenile stage. The specific gravity of foxtailed trees was found to be slightly less dense than that of normal trees though the differ-ence was not significant. Although breeding of true terminal foxtail trees may hold some promise of economic gains, the inherent limitations and foreseen problems render such proposition to be not feasible. Selection against foxtailing will continue to be a more practicable approach. Some future research studies on foxtailing are proposed: juvenile-mature correlation studies, long term growth and wood quality studies. i i TABLE OF CONTENTS Page ABSTRACT i TABLE OF CONTENTS i i LIST OF TABLES iv LIST OF FIGURES v ACKNOWLEDGEMENTS vi CHAPTER 1: INTRODUCTION 1 Section I - Objectives 2 Section II - Definition 2 CHAPTER 2: LITERATURE REVIEW 7 Section I - Foxtailing 7 A. Growth characteristics of foxtail trees ... 8 B. Occurrence of foxtail trees 9 C. Growth rate of foxtail trees 10 Section II - Specific Gravity 11 A. Variation in specific gravity 12 B. Specific gravity of foxtail trees 13 CHAPTER 3: MATERIALS AND METHODS 14 Section I - Description of study areas 14 Section II - Methods of sampling 17 A. Frequency and growth rate 17 B. Specific gravity 18 Section III - Counts and Measurements 20 A. Frequency count 20 B. Growth rate measurement 20 i i i Page C. Specific gravity determination 20 Section IV - Methods of Statistical Analysis .... 21 A. Frequency study 22 B. Growth rate study 23 C. Specific gravity study 24 CHAPTER 4: RESULTS AND DISCUSSION 27 Section I - Foxtail frequency 27 Section II - Growth rate 30 Section III - Specific gravity 32 Section IV - Prospects and limitations of breeding foxtail trees 34 CHAPTER 5: CONCLUSIONS 36 REFERENCES 38 APPENDIX I: Chi-square tests for proportion of foxtail to normal trees and proportion of sub-terminal to terminal foxtails 40 APPENDIX II: Analysis of variance for diameter over-bark of normal and foxtail trees 45 APPENDIX III: Analysis of variance for total height of normal and foxtail trees 50 APPENDIX IV: Analysis of variance for specific gravity of normal and foxtail trees 57 LIST OF TABLES iv Page CHAPTER 3: Table 3.1 Summary of c l i m a t i c data of Kemasul and Ulu Sedeli plantations 16 Table 3.2 Extent of 2 percent sampling i n Kemasul and Ulu Sedeli plantations 18 Table 3.3 Diameter over-bark and t o t a l height measure-ment of selected trees 19 Table 3.4 Chi-square test of binomial proportions 22 Table 3.5 Analysis of variance for diameter and t o t a l height of normal and f o x t a i l trees (sample of unequal s i z e s ) 23 Table 3.6 Analysis of variance for s p e c i f i c gravity ... 25 CHAPTER 4: Table 4.1 Frequency of f o x t a i l trees and proportion of various classes of f o x t a i l 28 Table 4.2 Mean diameters over-bark and t o t a l heights of normal and f o x t a i l trees 31 Table 4.3 Mean s p e c i f i c gravity of normal and f o x t a i l trees 32 V LIST OF FIGURES Page CHAPTER 1: Figure 1.1 Class 1A - True terminal foxtail 3 Figure 1.2 Class IB - Normal terminal foxtail 5 Figure 1.3 Class II - Sub-terminal foxtail 6 CHAPTER 3: Figure 3.1 Locations of Kemasul and Ulu Sedeli pine plantations and meteorological stations ... 15 CHAPTER 4: Figure 4.1 Effect of increasing height on specific gravity 33 ACKNOWLEDGEMENTS It is my pleasant duty to acknowledge my great indebtedness to the following: The Canadian International Development Agency for financing the study; the Forestry Department of Peninsular Malaysia for providing the materials and facilities used in this study; Professor Dr. 0. Sziklai and Dr. J.V. Thirgood of the Faculty of Forestry, U.B.C. and Dr. R.M. Kellogg of Forintek Canada Corp. for their constant guidance and encouragement; Mr. Ong Tai Hock, Research Assistant, Forest Research Institute, Peninsular Malaysia, for the field and laboratory assistance; and Robin Davison for reviewing the text. Lastly, but by no means least, I am most grateful to my wife, who relieved much of my domestic responsibility during the course of my absence. 1 CHAPTER I  INTRODUCTION Foxtailing, an extreme expression of apical dominance, is a common feature in plantations of Pinus caribaea var hondurensis Barrett and Golfari in Peninsular Malaysia and other tropical countries. This peculiar growth phenomenon is considered an undesirable trait. It is ' associated with wind breakage, development of compression wood, absence of late-wood formation and restriction of seed production (Kozlowski and Greathouse 1970; Wright 1976). However, foxtail trees often produce straight and knot-free timber. The narrow and conical-shaped crown of foxtail trees suggests that on a given planting area a higher volume production is expected from a pure "foxtail-tree" plantation than a pure "normal-tree" planta-tion. The narrow crown of foxtail trees permits higher stocking den-sity without causing any serious crown competition and early canopy closure. These characteristics give the impression that foxtailing may not be a totally undesirable trait. Kozlowski and Greathouse (1970) and Whyte et^  al. (1981) advocated breeding of foxtail trees for clean-boled trees. However, other traits of foxtail trees, such as growth rate and wood quality, need to be evaluated and compared with normal trees in order to make a valid appraisal of the value of foxtailing. A large portion of this study is directed at the investigation of frequency and growth rate of foxtail trees in Peninsular Malaysia. In addition a limited study on wood specific gravity in foxtail trees was carried out. 2 SECTION I - Objectives The objectives of this study are: 1. to determine the frequency of foxtail trees in plantations of Pinus caribaea var. hondurensis in Peninsular Malaysia, 2. to assess and compare the growth rate, in terms of diameter over-bark and height growth, of normal and foxtail trees, 3. to compare wood quality, in terms of specific gravity, of normal and foxtail trees, and 4. to evaluate the prospects and problems of breeding foxtail trees. SECTION II - Definition Foxtailing is an abnormal growth behaviour where a tree has a single, dominant and elongated shoot with no branches. In this study a stem which is longer than 0.8 metre with no branches is considered as foxtailed. There are two main classes of foxtail (Whyte er. al. , 1981), namely: Class I - Terminal foxtail, in which the stem continues to fox-tail from a certain point in the stem up to the tip of the tree. How-ever, in this study this class has been further sub-divided into two sub-classes; Class IA - True terminal foxtail, where continuous growth of the terminal leader results in a single stem with no side branches (Figure 1.1). Figure 1.1. Class IA - True terminal foxtail. A 4 year old true terminal foxtail with single dominant stem. Arrows show the formation of branches indicating that this true terminal foxtail is changing into a sub-terminal foxtail. 4 Class IB - Normal terminal f o x t a i l , where a normal pattern of shoot growth occurs during the e a r l i e r period and the f o x t a i l develops during l a t e r period (Figure 1.2). Class II - Sub-terminal f o x t a i l , i n which the top portion of the stem resumes a normal branched pattern a f t e r a period of f o x t a i l i n g growth (Figure 1.3). Figure 1.2. Class IB - Normal terminal foxtail. This 4 year old terminal foxtail developed after a normal shoot growth pattern. Figure 1.3. Class II - Sub-terminal foxtail. A 7 year old sub-terminal foxtail. Earlier foxtailing was followed by normal branch development. 7 CHAPTER 2 LITERATURE REVIEW Information available on foxtailing Is very limited. Lloyd (1914) was the first to describe the morphology of foxtailing in pine. Real interest in foxtailing began as recently as the early 1970's when Pinus caribaea Morelet gained recognition as one of the promising species for plantation forestry in many tropical countries. Foxtailing is a relatively new subject of research. To facilitate discussion of the available information this is considered under two sections: foxtailing and specific gravity. SECTION I - Foxtailing There is no difference in the pattern of normal shoot growth in the tropics and temperate zones (Kozlowski and Greathouse 1970). It involves the elongation of the axis by extension of a succession of buds formed on the terminal leader of the main stem. The period of elongation is interrupted for a while with the formation of new ter-minal bud clusters. These newly formed buds then expand to lengthen further the terminal leader and to produce a whorl of lateral branches. There may be two to four of such growth sequences annually. However, some abnormal growth of the tree may occur. In some pines, trees develop abnormally as a result of failure to set bud clusters which would differentiate into lateral branches. Such growth is known as "foxtailing" because the upper part of the abnormally elongated shoot has a conical or "fox-tail" appearance (Lloyd 1914). 8 There is no concrete evidence to explain what causes foxtailing. Based on the knowledge of bud dormancy and the balance of growth promoter-inhibitor in controlling shoot growth, Kozlowski and Greathouse (1970) speculated that in foxtailing growth there may be a continuous hormonal balance that favours one or more growth promoters which promote continuous shoot growth. Kozlowski and Greathouse (1970) also reported that foxtailing occurs in Pinus canariensis Smith, Pinus caribaea Morelet, Pinus  cembroides Link, Pinus echinata Link, Pinus e l l i o t t i i Engelm., Pinus  kesiya Royle ex Gordon, Pinus merkusii De Vriese, Pinus oocarpa Schiede, Pinus palustris Mill., Pinus radiata D. Don, Pinus taeda L. and Pinus tropicalis Morelet. A. Growth Characterisitics of Foxtail Trees In the shoot of a foxtail trees the needles usually decrease in length towards the terminal end. Near the tip, the unexpanded needles are tightly packed and enclosed in an unbroken sheath. Below these the needles penetrate their sheath and increase in length, giving the upper part of the continuous expanding shoot a conical appearance. Needle retention appears to be increased during the periods of foxtailing. This results in a distinctive dimpled bark pattern. The reproductive pattern of foxtail trees differs from that of normal trees (Kozlowski and Greathouse 1970). In a normal tree of Pinus caribaea, megasporangiate strobili are produced in the top fourth of the live crown while microsporangiate strobili are localised in the lower branches and on the third and fourth order side branches. During the periods of apical foxtail growth no strobili were reported on trees 9 up to 15 years old. However, the terminal shoots of foxtail trees are said occasionally to produce a large number of microsporangiate strobili. B. Occurrence of Foxtail Trees Among the tropical pines, such as Pinus caribaea, Pinus oocarpa and Pinus merkusii, which have been planted in Peninsular Malaysia, foxtailing is more frequently observed in Pinus caribaea. Tho (1979) reported that among the three species mentioned above, there was 31.7 percent foxtailing in Pinus caribaea, 19.5 percent in Pinus merkusii and 0 percent in Pinus oocarpa. Within the three varieties of Pinus  caribaea namely var. hondurensis Barrett and Golfari, var. bahamensis Barrett and Golfari and var. caribaea Barrett and Golfari, foxtailing is common in var. hondurensis (31.7 percent) as compared to 8.0 percent in var. bahamensis and 0 percent in var. caribaea (Musalem et_ al., 1973; Wiersum 1973). Foxtailing of Pinus caribaea var, hondurensis appears to be an inherited growth phenomenon with its expression modified considerably by site and climatic factors. In an experiment of family variation within provenances for foxtailing growth form in Pinus caribaea var. hondurensis, Ledig and Whitmore (1980) reported 0.17 heritability of foxtailing. Although this value is low, selection against foxtailing could s t i l l be effective. Earlier in 1968, Slee and Nikles indicated that an unselected stand of Pinus caribaea var. hondurensis in Queens-land, Australia, had a much higher frequency of foxtail than did unselected var. bahamensis and var. caribaea. However, after selection the progenies of selected var. hondurensis parents had a lower incidence of foxtailing. 10 The climatic control of foxtailing also appears to be pronounced. Lucknoff in 1964 (quoted by Kozlowski and Greathouse 1970) observed that foxtailing of Pinus caribaea was reduced at higher elevation and cooler temperature. In coastal areas of Zululand (altitude approxi-mately 45 to 60 metres) foxtail frequency was 43 percent; at Ntsubane (altitude approximately 400 metres) 26 percent; and at Dargal in the Natal Midlands (altitude approximately 1200 metres) 13 percent. Occurrence of foxtailing also varies with site quality. Using a single provenance of Pinus caribaea var. hondurensis seeds, Ibrahim and Greathouse (1972) observed different frequencies of foxtail when the seeds were planted at two different locations in Peninsular Malaysia. On the fertile site the foxtail frequency was 39 percent while on the poorer site the frequency was 49 percent. C. Growth Rate of Foxtail Trees Except for the peculiar crown of foxtail trees, their growth performance, in terms of diameter and height, is comparable and at times superior to normal trees. Most reports (Ibrahim and Greathouse 1972; Wood et al., 1979 and Whyte et_ al., 1981) indicate that foxtail trees have superior height growth compared to normal trees. However, the diameter growth of foxtail trees varies with age as compared to normal trees. In a 6 year old plantation foxtail trees have smaller diameter than normal trees (Ibrahim and Greathouse 1972). Similarly, Wood et al. (1979) also indicated that in a 6 year old plantation the average diameter of foxtail trees was significantly less than normal trees. In a frequency 11 and growth rate study of foxtail trees in plantations aged 3 to 9 years, Whyte et al. (1981) concluded that foxtail trees were larger in diameter up to age 5 years but smaller thereafter when compared to normal trees. SECTION II - Specific Gravity Specific gravity is defined as the ratio of the density of a given substance to the density of water at standard temperature and pressure. The specific gravity of wood may be based on volume when oven dry (dried at 105°C to constant bone-dry weight); or when green (at fully swollen condition, at some moisture content above the fibre saturation point); or at some moisture content intermediate between these two extreme conditions. Due to the probability of unequal shrinkage in different wood samples when drying from green to oven dry weight, specific gravity based on oven dry weight and green volume is the preferred measure in all cases when comparisons are made (Smith 1954). Due to the ease of measurement and straightforward interpreta-tion, specific gravity is an excellent index of the amount of wood substance contained in a given volume of wood. Therefore it is a good indicator of strength properties (U.S.D.A. 1955). There is a strong correlation between wood specific gravity and compression, bending strength and hardness (Panshin and De Zeeuw 1980). Specific gravity and wood density are usually used synonymously when the former is based on oven dry weight and green volume and the latter is measured in grams per cubic centimetre green volume. 12 A. V a r i a t i o n i n S p e c i f i c Gravity A piece of dry timber i s composed of s o l i d material ( c e l l walls) and c e l l c a v i t i e s . The difference i n the r a t i o of c e l l walls to c e l l c a v i t i e s gives r i s e to the difference i n wood s p e c i f i c gravity within and between trees (Desch 1981). Variations i n the amount of c e l l wall substance i n wood are due to the changes of the anatomical characteris-t i c s of the c e l l wall and proportion of d i f f e r e n t c e l l types occurring i n d i f f e r e n t parts of the trees. There are b a s i c a l l y two general trends of v a r i a b i l i t y i n s p e c i f i c gravity within the tree, namely v a r i a t i o n of s p e c i f i c gravity along the stem length and v a r i a b i l i t y of s p e c i f i c gravity i n the stem cross section ( E l l i o t t 1970; Panshin and De Zeeuw 1980). In many coniferous species the heaviest wood i s found at the base of the trunk and decreases i n successively higher l e v e l s i n the trunk, and at any given height of the tree trunk s p e c i f i c gravity increases from the p i t h out-wards. The former v a r i a t i o n i s due to the increase percentage of young material in the successive increment from the base to the top of the trunk and the l a t t e r i s due to the increased proportion of dense l a t e -wood i n the successive increment from the p i t h outwards. Harris (1973) observed a steep density gradient from the p i t h outwards i n Pinus caribaea grown i n Peninsular Malaysia- I t appeared that the increase in wood density from the p i t h outwards terminated between the 8th and 12th growth layer. M u l t i p l e bands of very dense late-wood i n each annual growth layer were also observed. 13 The variation in specific gravity among trees of the same species is influenced by inherent characteristics of the tree, geographical and environmental factors (Harris 1978; Panshin and De Zeeuw 1980; Cown 1981). Silvicultural practices, such as fertilizer application and manipulation of crown size and growing space to promote growth rate, have significant influence on specific gravity (Elliott 1970 and Panshin and De Zeeuw 1980). B. Specific Gravity of Foxtail Trees It appears that the specific gravity of foxtail trees is lower than that of normal trees. Plumptre (1978) and Whyte et al. (1981) made comparative studies of specific gravity between normal and foxtail trees of Pinus caribaea using samples from an 8 year old plantation. Both reports indicated that timber from foxtail trees was less dense than that of normal trees by 8 percent (Plumptre 1978) and by 0.8 per-cent (Whyte et al., 1981). 14 CHAPTER 3 MATERIALS AND METHODS This chapter is divided into four main sections: description of study area; methods of sampling; frequency count, growth rate measure-ment and specific gravity determination; and methods of statistical analysis. Section I - Description of Study area This study was carried out in two Pinus caribaea var. hondurensis plantations in Peninsular Malaysia, namely Kemasul Pine Plantation and Ulu Sedeli Pine Plantation (Figure 3.1). The Kemasul plantation is located about 145 kilometres east of Kuala Lumpur, covering an area of 25,000 hectares of which 22,000 hectares are suitable for reforestation (Anon. 1983). The Ulu Sedeli plantation is situated about 400 kilometres southeast of Kuala Lumpur, covering an area of 34,885 hectares of which 29,434 hectares are suitable for planting (Anon. 1982). Both plantations are in the undulating lowland areas with an average altitude of 60 metres above sea level. Table 3.1 summarized the climatic data of both plantations for five year period, 1977-1981. In this period, it seems that both plantations appeared to have, similar climatic conditions though Ulu Sedeli plantation is slightly wetter than Kemasul plantation. The soil varies between and within sites. The most common soil in Kemasul is derived from sandstone and shale (Teoh 1981), while in 15 Figure 3.1 Location of Kemasul and Ulu Sedeli pine plantations i n Peninsular Malaysia. 16 TABLE 3.1 Summary of climatic data of Kemasul and Ulu Sedeli plantations Meteorological Year Station Max. Daily Temperature (°C) Min. Daily Temperature <°C) Total Annual Rainfall (mm) Bentong* 1977 1978 1979 1980 1981 Average 32.2 33.4 34.1 33.1 32.9 33.3 22.4 21.4 21.2 22.1 22.1 21.9 2280.4 1747.9 2560.0 na 2098.4 2171.7 Kota Tinggi** Average 1977 1978 1979 1980 1981 31.9 32.2 32.3 32.2 31.6 32.0 22.0 21.8 21.8 22.7 22.8 22.2 2118.8 2501.3 2505.0 2359.7 1999.0 2296.0 * and ** - Meteorological stations nearest to Kemasul and Ulu Sedeli plantations, respectively (Figure 3.1). na - Not available. Ulu Sedeli the soils are mostly derived from granodiorite (Amir 1983, personal communication). Planting in both sites started in early 1975. To date about 2700 hectares have been planted in Kemasul and about 1750 hectares in Ulu Sedeli. No planting was carried out in 1980 in Kemasul and in 1982 in Ulu Sedeli. The planting distance used in both plantations is 9 feet by 7 feet (2.7 metres by 2.1 metres), along the east-west direction, 17 with a stocking density of 691 trees per acre (1780 trees per hectare). Seedlings were raised from seeds purchased overseas. From 1974 to 1976 seeds were obtained from Central America. After the seed supply from Central America was discontinued, Australia became the seed supplier in 1977. Due to the high cost and limited seed supply for Australia, the F i j i Pine Commission became the current seed supplier since 1979 t i l l today. SECTION II - Methods of Sampling A. Frequency and Growth Rate Studies In both plantations the annual planting areas are normally divided into several blocks which range from 10 to 100 hectares. In view of this fact a stratified random sampling without replacement was used in this study. In each plantation the population was stratified into 7 age classes. From each age class a planting block was randomly selected. In the selected block a linear sampling of 2 percent inten-sity was carried out to determine the frequency and growth rate (dia-meter over-bark and total height) of normal and foxtail trees. Table 3.2 indicates the extent of sampling carried out. A sampling line consists of 4 rows of trees which run across the block along the east-west direction. In determining the initial sampling line, a point was selected randomly from a set of distances measured from the left-hand corner of the block: 5, 10, 15, 20 and 25 metres. In this case a distance of 10 metres was selected and used to set the initial sampling line throughout sampling of the entire popula-tion of the plantation. 18 TABLE 3.2 Extent of 2 percent sampling in Kemasul and Ulu Sedeli plantations Plantation Age Selected Area of selected Area No of trees (year) block block (ha) sampled (ha) sampled 1 2C3 62.8 1.25 1196 2 2A5 56.7 1.13 1127 4 ID 5 57.0 1.14 448 Kemasul 5 1D6 68.9 1.37 756 6 1C7 32.0 0.64 640 7 1B4 44.5 0.89 416 8 1A3 30.4 0.60 854 Total 352.3 7.02 5437 2 5.7 22.3 0.44 304 3 4.11 40.4 0.80 511 4 4.6 30.7 0.61 339 Ulu Sedeli 5 4.1 47.3 0.94 670 6 3.4C 18.0 0.36 302 7 2.2C 38.4 0.76 559 8 1.2 37.3 0.74 816 Total 234.4 4.65 3501 Depending on the size and shape of the block one or more sampling lines were required to cover the 2 percent sampling intensity. In blocks where more than one sampling line was needed the subsequent lines were set at 10 metre intervals from the previous line. B. Specific Gravity A complete randomised design with hierarchal arrangement was used in this study. In each plantation five normal trees and five foxtail trees were selected at random from the 8 year-old stand. Table 3.3 indicates the size of the selected trees. 19 TABLE 3.3 Diameter over-bark and total height measurements of selected trees Normal Foxtail* Plantation Diameter (cm) Total Height (m) Diameter (cm) Total Height (m) 10.5 10.3 12.7 12.1 10.5 10.3 11.7 12.1 Kemasul 9.8 10.3 11.7 10.9 11.6 10.6 8.2 10.3 11.4 9.1 10.5 10.3 13.2 13.7 12.5 18.2 13.2 12.8 11.3 18.2 Ulu Sedeli 13.5 12.1 11.3 15.3 12.7 12.1 11.0 14.3 11.8 12.1 11.6 15.2 *Foxtail of Class I where foxtail occurs from 10 percent of total height upwards. A disc of 2 centimetres in thickness was taken from each tree at five different percentage-height levels of the tree: 10; 30; 50; 70; 90 percent. Each disc was divided into four quarters as replications. The samples were then debarked and prepared for testing. Percentage-height sampling is chosen over sampling at fixed position along the stem because it allows for between tree comparison and at the same time accounts for the systematic variation of specific gravity within the stem (Elliott 1970). 20 SECTION III - Frequency Count, Growth Rate Measurement and Specific  Gravity Determination A- Frequency Count In the sampling line the occurrence of normal and foxtail trees was recorded and tallied. For foxtail trees, the trees were further categorised according to their classes. B. Growth Rate Measurement Except in the 1 and 2 year old age classes diameter over-bark at breast height (1.4 metres) of normal and foxtail trees was measured using a diameter tape (measured to the nearest 0.1 centimetre). Height sticks were used to measure the total height of normal and foxtail trees in all age classes (measured to the nearest 0.1 metre). In addi-tion, measurement of the longest internode length (length between 2 whorls of branches) of normal and foxtail trees in the 4, 7 and 8 year old stand at Kemasul were also taken. These measurements were used to define foxtail characteristics quantitatively as mentioned in Chapter I. C• Specific Gravity Determination Specific gravity is expressed as the ratio of the density of a given substance to that of water. Specific gravity is therefore a unit-less value but numerically equal to the density of the substance. In this study specific gravity or density was determined gravimetri-cally based on the oven dry weight and green volume of sample. The formula used for wood density is: , , . Oven dry weight of sample Wood density = — —i—-— — Green volume of sample 21 The green volume of the sample was determined as follows: A beaker of water was placed on a balance pre-set to the weight of the beaker and water. The test sample suspended by a needle was lowered in the beaker and completely immersed in water. Care was taken so that the immersed sample was not in contact with either the sides or bottom of the beaker. The reading on the balance was then recorded. The weight of the displaced volume of water represents the green volume of the sample. The procedure to determine the oven dry weight of the sample was as follows: The labelled samples were placed in a controlled temperature oven at 105°C for 48 hours. At the end of the period 15 samples were chosen at random and placed in a desiccator containing granulated anhydrous calcium chloride (CaCl2.2H.2O). After a period of conditioning at laboratory temperature the samples were weighed on a balance (weighed to 3 decimal places). After the first oven dry weight was determined, samples were replaced in the oven for another 24 hours under the same temperature condition. This cycle was repeated until a constant oven dry weight was recorded. SECTION IV - Methods of Statistical Analysis Two standard statistical texts were used as reference sources for the analyses computed in this study: Spiegel (1972) and Snedecor and Cochran (1980). 22 A. Frequency Study This study deals with two classes of individuals in a population: normal and foxtail trees. Since a tree can either be normal or foxtail in form, the outcomes of this study can be expressed as percentage, proportion or number of individuals in one of the two classes. These conditions fit the binomial distribution model. Chi-square test for goodness of f i t in a RxC contingency table is used to compare the proportions of normal to foxtail trees and sub-terminal and terminal foxtails. The hypotheses tested are: the pro-portion of normal to foxtail trees is constant over age; and proportion of sub-terminal to terminal foxtail is constant over age. Before carrying out the test the proportions are transformed by angular trans-formation (arc-sine). The test is computed in the following manner (Table 3.4): TABLE 3.4 Chi-square test of binomial proportions Component Element I Element II Observed, f r n-r Expected, F np nq = n-np Observed-Expected, f-F r-np -(r-np) 2 - .(f-F) _ (r - np)2 (r - np)2 Y — h _ — i "T • • • • F np np where: Element I = either normal tree or sub-terminal foxtail Element II = either foxtail tree or terminal foxtail 23 r observed number of Element I n total number of sample P proportion of Element I q proportion of Element II Degree of freedom = (R-l) (C-l) where: R = rows C - Columns B. Growth Rate Study Analysis of variance is used to compare the mean of diameter and total height between normal and foxtail trees within each age class and plantation. The analysis is based on unequal samples and carried out in the following manner (Table 3.5): TABLE 3.5 Analysis of variance for diameter and total height of normal and foxtail trees (samples of unequal sizes) Source of variation df SS MS F Between phenotypes a-1 SSI MSI MSI/MSII Within phenotypes N-a SSII MSII Total N-l where: a = phenotypes (normal and foxtail trees) N = number of samples 24 C. S p e c i f i c Gravity Study The mean s p e c i f i c gravity of normal and f o x t a i l trees was com-pared by a one-way analysis of variance. The analysis was car r i e d out for each percentage-height l e v e l separately. This study was designed i n a completely randomised manner involving h i e r a r c h a l arrangement. The sources of v a r i a t i o n , degrees of freedom and expected mean squares for the analysis of variance are shown i n Table 3.6. The F - r a t i o s were determined as follows: MSIII ( i ) Between trees/phenotypes/plantations: F = MS IV where: MSIII = mean square for between trees/phenotypes/ plantations MSIV = mean square for r e s i d u a l . ( i i ) I f F - r a t i o ( i ) i s s i g n i f i c a n t then F - r a t i o for between phenotypes/plantations i s : F = MSII MSIII where: MSII = mean square for between phenotypes/ plantations ( i i i ) I f F - r a t i o ( i i ) i s not s i g n i f i c a n t then a new mean square error i s derived to test the v a r i a t i o n between plantations: SSII + SSIII MSE = df2 + df3 where: MSE' = new mean square error SSII = sum square between phenotypes/plantations SSIII = sum square between trees/phenotypes/ plantations TABLE 3.6 Analysis of variance for specific gravity Source of variation df SS MS Expected mean square Between plantations ( P L ) dfl = (a-1) SSI MSI a 2 + K L A ^ / P H / P L + V ^ H / P L + S ^ P L Between phenotypes/ df2 = a(b-l) SSII MSII a2 + \oZ . . + K ^ P H / P L plantations ( P H / P L ) 2 2 Between trees/phenotypes/ df3 = ab(c-l) SSIII MSIII a + K CT T / M , / B T plantations ( T / P H / P L ) 6 1 T / P H / P L Residual df4 = abc(n-l) SSIV MSIV a 2 e Total abcn-1 where: a = plantations b = phenotypes c = trees n = number of samples to K3 = coefficients of the variance components. df2 = degrees of freedom between phenotyp< plantations df3 = degrees of freedom between trees/phenotypes/ plantations Then, F-ratio for between plantations: F = ^|^r, where: MSI = mean square between plantations. 27 CHAPTER 4 RESULTS AND DISCUSSION The presentation of this chapter is divided into four main sections: foxtail frequency; growth rate; specific gravity; and prospects and limitations of breeding foxtail trees. SECTION I - Foxtail Frequency The study shows that the frequency of foxtail at Kemasul planta-tions aged 1 to 8 years varies between 6.1 to 36.0 percent, while at Ulu Sedeli plantations aged 2 to 8 years the frequency varies between 4.2 to 34.8 percent (Table 4.1). The overall foxtail frequencies at Kemasul and Ulu Sedeli are 18.1 + 1.0 percent and 23.4 + 1.4 percent respectively, a difference of 5.3 percent. More than 60 percent of foxtails in both plantations are in the form of sub-terminal foxtail: 70.8 and 64.2 percent in Kemasul and Ulu Sedeli respectively. Table 4.1 also indicates that the proportions of foxtail to normal trees and sub-terminal and terminal foxtails tend to increase with age. The Chi-square tests (Appendix I - Tables 1 to 4) confirm that these proportions are not the same throughout the age classes. The difference in foxtail frequency within each plantation observed in this study is suspected to be due to the differences in seed source, site and age. This observation conforms to the observa-tions made by Ibrahim and Greathouse (1972) and Musalem et al. (1973). Between plantations the difference in foxtail frequency, 5.3 percent 28 TABLE 4.1 Frequency of foxtail trees and proportion of various classes of foxtail (%) Plantation Age Foxtail Sub-terminal Terminal Terminal foxtail (year) foxtail foxtail Normal True 1 6.9 52.4 47.6 75.6 24.4 2 6.1 71.0 29.0 80.0 20.0 4 19.2 51.0 49.0 87.7 2.3 Kemasul 5 19.7 41.7 58.3 100 0 6 25.4 89.0 11.0 100 0 7 36.0 96.7 3.3 100 0 8 33.3 94.1 6.0 94.1 5.9 Average foxtail frequency 18.1 +1.0 2 4.2 46.2 53.8 100 0 3 11.4 51.7 48.3 96.4 3.6 4 8.5 55.2 44.8 100 0 Ulu Sedeli 5 23.7 57.2 42.8 100 0 6 34.8 67.6 32.4 94.1 5.9 7 31.1 87.4 12.6 100 0 8 34.4 84.0 16.0 95.4 4.4 Average foxtail frequency 23.4 +1-3 as indicated in this study, is probably due to site quality; higher frequency in poorer site (Slee and Nikles 1968; Ibrahim and Greathouse 1972). The Kemasul plantation is a better site than Ulu Sedeli planta-tion based on growth rate indicated in Section II of this chapter. The design of this study does not permit the investigator to verify the speculation that proportions of foxtail to normal trees and sub-terminal to terminal foxtails increase with age. This may require the assessment and observation of the same population over a certain period of time in order to ascertain the trend of the changes. How-ever, the work of Ibrahim and Greathouse (1972) indicates that foxtail 29 frequency increases with age. Observing the same population of Pinus  caribaea var. hondurensis, raised from seeds of a single provenance, over a period of three years the foxtail frequency increases by almost three times. The work of Whyte et al_. (1981) indicates that the pro-portion of sub-terminal foxtail increases from 5 to 12 percent and the proportion of terminal foxtail decreases from 19 to 8 percent in a 3 year old plantation two years after the first assessment. This explains the higher frequency of foxtail trees and sub-terminal fox-tails observed in the older stands of the study sites. The increasing proportion of sub-terminal foxtails over time suggests that foxtailing is a plastic trait, where it changes its form from terminal to sub-terminal . All foxtail trees start in the form of terminal foxtail. In the course of its growth, a foxtail tree may either continue to show extreme apical dominance or rest to set lateral buds to produce side branches. Once the foxtail tree produces lateral branches the form changes from terminal to sub-terminal foxtail. This may mark the end of foxtailing growth or temporary conversion to normal growth pattern before foxtailing growth occurs again in later years. This means that foxtail is an unstable trait. The foxtail frequency study also shows among the terminal foxtails the occurrence of true terminal foxtail is very low compared to normal foxtail. It is probable that the tendency for the true terminal foxtail to maintain its form is lowered with increasing age of the tree. 30 SECTION II - Growth Rate Table 4.2 indicates that the overall growth of trees in Kemasul plantation is superior to that in Ulu Sedeli plantation especially in diameter growth. This may suggest that the Kemasul plantation is a better site than the Ulu Sedeli plantation. The table also indicates that the diameter growth of foxtail trees surpasses the normal trees in the early years of growth from age 4 to 5 years and 3 to 7 years in Kemasul and Ulu Sedeli plantations respectively. The analysis of variance (Appendix II - Tables 1 to 11) shows that the superiority of foxtail trees in diameter growth over normal trees is significant only in 4 year old trees at Kemasul plantation and in 3, 4 and 5 year old trees at Ulu Sedeli plantation. In terms of height growth, foxtail trees maintain their supremacy up to 6 years old in Kemasul and 8 years old in Ulu Sedeli (Table 4.2). However, this supremacy is only significant to year three and year four at Ulu Sedeli and Kemasul plantations respectively (Appendix III -Tables 1 to 14). Generally, the results indicate that foxtail trees are signifi-cantly superior to normal trees in diameter and height growth especially in the first 4 to 5 years of growth. These findings confirm the observations made by Ibrahim and Greathouse (1972), Wood et al. (1979) and Whyte et al. (1981). 31 TABLE 4.2 Mean diameters over-bark and total heights of normal and foxtail trees Plantation Age (year) Diameter over-bark (cm) Total height (m) Normal Foxtail F-test Normal Foxtail F-test 1 0.9 1.2 ft* 2 - - - 1.8 2.3 ft** 4 9.7 10.1 * 6.1 7.0 *** Kemasul 5 8.9 9.3 ns 5.6 6.6 ns 6 12.8 12.7 ns 9.6 10.2 ns 7 17.6 16.6 *** 11.6 11.5 ns 8 13.7 13.4 ns 10.6 11.2 ns 2 1.8 3.2 *** 3 2.5 3.9 ** 3.3 4.4 * 4 2.7 5.7 *** 3.8 3.9 ns Ulu Sedeli 5 3.9 5.8 *** 6.4 6.9 ns 6 5.4 6.5 ns 7.9 8.5 ns 7 4.7 4.8 ns 9.3 9.4 ns 8 7.1 6.1 ns 11.2 12.2 ns These results also suggest that most of the foxtailing growth phenomenon In a population occurs between 1 to 4 years. This is indicated by the rapid height growth as a result of extreme apical dominance shown by foxtailing growth. After such a period most of the terminal foxtails become sub-terminal foxtails as discussed in the last section (Section I), and lose their vigour in height growth. This probably explains the non-significant height growth between normal and foxtail trees after 4 years of age. 32 SECTION III - Specific Gravity Table 4.3 represents the mean of specific gravity of normal and foxtail trees sampled at five different percentage-height levels in Kemasul and Ulu Sedeli plantations. Except for the 10 percent height TABLE 4.3 Mean specific gravity of normal and foxtail trees Height level (%) Normal Foxtail F-test 10 0.542 0.569 ns 30 0.519 0.499 ns 50 0.490 0.460 ns 70 0.453 0.414 ns 90 0.404 0.362 ns level, foxtail trees are slightly less dense than normal trees. The difference in mean specific gravity between normal and foxtail trees at all height levels was found to be not significant (Appendix IV - Tables 1 to 5). The finding of this study agrees with the observations made by Plumptre (1978) and Whyte et al. (1981) that the specific gravity of foxtail trees is slightly lower than normal trees. However, the difference is highly significant between trees within phenotypes and plantations. This means that variation in specific gravity is only due to variation among trees with the phenotype. The result also indicates that there is an obvious pattern in both normal and foxtail trees, that specific gravity decreases with increasing height (Figure 4.1). 34 SECTION IV - Prospects and Limitations of Breeding Foxtail Trees This study indicates that foxtail trees have fast early diameter growth, superior height growth and a specific gravity slightly less dense than normal trees. These characteristics coupled with a generally straight stem form (Whyte et_ al., 1981) and obvious narrow crown especially with the true terminal foxtail trees suggest that breeding of true terminal foxtail trees may hold promise of some economic gains. Plantations of true terminal foxtail tree can be envisaged as pure stands with fast early growth producing a uniform product of knot-free timber. Maximum utilization of growing space is visualized as true terminal foxtail trees can be planted at a closer spacing without experiencing serious canopy competition and early canopy closure. It is also foreseen that harvesting technique and log transportation from such plantations will be simpler and more economical than the same operations in heterogenous plantations. However, there are some inherent limitations and foreseen prob-lems to such breeding programme. As has been discussed earlier, true terminal foxtail is a very plastic or unstable form. Its occurrence is quite scarce and its ability to maintain such form throughout the rotation age is s t i l l questionable. This poses a major set back in initiating a breeding programme for such trait. Also its heritability is low (Ledig and Whitmore 1981), therefore a high selection intensity is required to obtain the desired genetic gain. This also means that a large breeding population is needed. Evidence from Peninsular Malaysia indicated that locally grown Pinus caribaea is known to be a shy-seeder (Mitchell 1963; Freezaillah 35 1967). Therefore, the probability for foxtail trees to produce seeds in Peninsular Malaysia is very low if not nil. Furthermore, based on the observations of Kozlowski and Greathouse (1970) foxtail trees were not known to produce seeds. This will certainly cause major problem if the plantations of true terminal foxtail trees are to be raised from seeds. Mass propagation by stem cuttings of true terminal foxtail trees is impossible because of the absence of branches to obtain the cuttings. The other alternative is propagation by rooting needle fascicles or by means of tissue culture. Neither technique has yet been proven successful in Pinus caribaea. Foxtailed trees often exhibit an above average incidence of wind breakage, compression wood formation and juvenile wood (Kozlowski and Greathouse 1970; Wright 1976). The presence of compression wood and juvenile wood will substantially reduce the quality of the timber. The amount of compression wood is influenced by the interaction of tree lean, growth rate and slope of terrain. The true terminal fox-tail tree which is normally tall and branchless in form creates a very unstable structure which is susceptible to lean, sway or bend by exter-nal forces such as wind. This may cause the tree to produce compres-sion wood in order to restore the leaning tree stem to its normal ver-tical orientation. Juvenile wood is formed about the pith as a result of prolonged influence of apical meristems in the region of active crown on wood formation by the cambium (Panshin and De Zeeuw 1980). The long and vigorous crown of foxtail trees will favour the formation of juvenile wood rather than mature wood. 36 CHAPTER 5  CONCLUSIONS Based on the study carried out during the summer of 1983, the frequency of foxtail trees, in the plantations of Pinus caribaea var hondurensis in Peninsular Malaysia, varies between and within planta-tions. In Kemasul pine plantation an overall foxtail frequency is 18.1 +1.0 percent and it ranges from 6.1 to 36.0 percent, while in Ulu Sedeli pine plantation the frequency ranges from 4.2 to 34.8 percent with an overall frequency of 23.4 + 1.3 percent. This study indicates that the foxtail frequency increases with age, however, there is no evidence yet to determine at what age will the frequency stabilize. It is also found that the overall foxtail frequency in Ulu Sedeli plantation is higher than in Kemasul plantation by 5.3 percent. This further indicates that foxtail frequency also varies with site. The most common form of foxtail in both plantations is the sub-terminal foxtail (more than 60 percent) while the occurrence of true terminal foxtail is very low. It appears that the proportion of sub-terminal foxtail to terminal foxtail increases with age which suggests that foxtailing is a plastic trait. In terms of diameter growth, the superiority of foxtail trees is evident only during the early stages of growth. Generally, foxtail trees are superior in height growth than normal trees, however the superiority in height growth is significant in the first four years of growth. 37 In wood specific gravity, foxtail trees are slightly less dense than normal trees. However, the difference is not significant. Both normal and foxtail trees exhibit the systematic pattern of variation in which specific gravity decreases with increasing height. Although foxtail trees, especially the true terminal foxtail, hold promise of some economic gains, the inherent limitations and fore-seen problems suggest that breeding of foxtail trees may not be feasible and practicable. This is due to the plasticity of the trait, restriction of seed production and formation of juvenile and compres-sion wood. Taking advantage of the low heritability of foxtailing trait, selection against foxtail could be very effective in environ-ments that highly favour the incidence of foxtailing. Much of the information of growth characteristics of foxtail trees are based on short term observations. Long term observations will provide more reliable information as to the changes of growth of foxtail trees over time. Wood quality studies of foxtail trees in relation to compression and juvenile wood formation are needed to explain and determine the extent of juvenile and compression wood formation in foxtail trees. Particularly in Peninsular Malaysia, where it has to rely upon overseas seed supply, foxtailing will s t i l l be a common feature in the local pine plantations. Selection against foxtailing could be more effective at seed source and seedling levels. Seed and juvenile-mature correlation studies of the foxtailing trait could provide useful guidelines for early selection against foxtailing. 38 REFERENCES Anon- 1982. Progress report on pine plantation in Ulu Sedeli Forest Reserve, Kota Tinggi, Johor. 10 pp. (Text in Malay). Anon. 1983. Report on pine plantations in Kemasul, Pahang. 4 pp. (Text in Malay). Cown, D.J. 1981. Wood density of Pinus caribaea var. hondurensis grown in F i j i . New Zealand Journal of Forestry 11(3).244-253. Desch, H.E. 1981. Timber: Its structure, properties and utilization. 6th ed. Timber Press, Oregon. 410 pp. Elliott, G.K. 1970. Wood density in conifers. Technical Communica-tion No. 8. Commonwealth Agricultural Bureau, England. 44 pp. Freezaillah, C.Y. 1973. Pilot plantation for quick-growing industrial tree species. Malaysian Forester 30(4):246-252. Harris, J.M. 1973. The use of beta rays to examine wood density of tropical pine grown in Malaya. In: Burley, J. and Nikles, D.G. (eds.), Selection and Breeding to Improve Some Tropical Conifers. Vol. 2. Commonwealth Forestry Institute, Oxford: 86-94. Harris, J.M. 1978. Note on wood density of Pinus caribaea Morelet grown under temperate, sub-tropical and tropical conditions. In: Nikles, D.G., Burley, J. and Barnes, R.D. (eds.), Progress and Problems of Genetic Improvement of Tropical Forest Trees. Vol. 1. Commonwealth Forestry Institute, Oxford: 1-7. Ibrahim, S. and T.E. Greathouse. 1972. Foxtailing in exotic pine -preliminary results of study in West Malaysia. Malaysian Forester 35(l):17-23. Kozlowski, T.T. and T.E. Greathouse. 1970. Shoot growth and form of pines in the tropics. Unasylva 24(4):6-14. Ledig, F.T. and J.L. Whitmore. 1981. Heritability and genetic correlations for volume, foxtails and other characteristics of Caribbean pine in Puerto Rico. Silvae Genetic 30(2-3):88-92. Lloyd, F.E. 1914. Morphological instability especially in Pinus  radiata. Botanical Gazette 57:314-319. Lucknoff, H.A. 1964. The natural distribution, growth and botanical variation of Pinus caribaea Mor. and its cultivation in South Africa. Ann. Univ. Stellenbosch 39:1-160. Mitchell, B.A. 1963. Possibilities for forest plantations. Malaysian Forester 26(4):259-286. 39 Musalem, M.A. and P. Rozero. 1973. Performance of varieties and provenance of Pinus caribaea introduced in Turrialba, Costa Rica. Turrialba 23(3):327-333. Panshin, A.J. and C. De Zeeuw. 1980. Textbook of Wood Technology. 4th ed. McGraw H i l l , New York. 665 pp. Plumptre, R.A. 1978. Variation in wood density of Pinus caribaea var. hondurensis Barrett and Golfari grown in plantation. Method developed to study i t . In: Nikles, D.G., Burley, J. and Barnes, R.D. (eds.), Progress and Problems of Genetic Improvement of Tropical Forest Trees. Vol. 1. Commonwealth Forestry Institute, Oxford: 46-45. Slee, M.U. and D.G. Nikles. 1968. Variability of Pinus caribaea Mor. in young Queensland plantations. Proceeding of the 9th Common-wealth Forestry Conference, New Delhi: 1-50. Smith, D.M. 1954. Maximum moisture content method for determining specific gravity of small wood samples. Report 2014, U.S.D.A. Forest Product Laboratory, Madison. Snedecor, G.W. and W.G. Cochran. 1980. Statistical Methods. 7th ed. Iowa State University Press, Iowa. 459 pp. Spiegel, M.R. 1972. Theory and Problems of Statistics. McGraw H i l l , New York. 339 pp. Tho, Y.P. 1979. A note of foxtailing and multiple leaders in exotic pine at Bahau experimental pine plantations. Malaysian Forester 42(3):255-277. Teoh, S.K. 1981. Soil suitability in relation to Caribbean pine growth and yield. Malaysian Forester 44(1):60-73. U.S.D.A. 1955. Wood Handbook. U.S.D.A. Forest Product Laboratory, Madison. Whyte, A.G.D., P.M. Adams and S.E. McEwen. 1981. Foxtailing of Pinus  caribaea var. hondurensis in F i j i : frequency, distribution of occurrence and wood properties. Forestry Ecology and Management 2(3):237-243. Wiersum, K.F. 1973. Some observation of the occurrence of foxtails in young Pinus caribaea plantation at Turrialba. Turrialba 23(3):362-365. Wood, F.W., L.W. Vincent, W.W. Moscheler, and H.A. Core. 1979. Height, diameter, and specific gravity of 'foxtail' trees of Pinus caribaea Mor. Forest Product Journal 29(5):43-44. Wright, J.W. 1976. Introduction to Forest Genetics. Academic Press, New York. 436 pp. APPENDIX I Chi-square Test - Proportion of foxtail to normal trees and proportion of sub-terminal to terminal foxtails Legend * - significant. 5% level of probability ** - significant. 1% level of probability *** - significant. 0.1% level of probability ns - not significant. 41 TABLE 1. Chi-square test of the proportions of foxtail to normal trees at Kemasul plantation Age (year) 8 Total Frequency observed, f Normal 74.7 75.7 64.0 63.6 59.7 53.1 54.7 445.5 Foxtail 15.3 14.3 26.0 26.4 30.3 36.9 35.3 184.5 Total 90.0 90.0 90.0 90.0 90.0 90.0 90.0 630.0 Frequency expected, F Normal Foxtail 63.6 26.4 63.6 26.4 63.6 26.4 63.6 26.4 63.6 26.4 63.6 26.4 63.6 26.4 (f-F) Normal Foxtail 1.94 4.67 2.30 5.55 0.002 0.00 0.006 0.00 0.24 0.58 1.73 1.24 4.18 3.00 2 _ ,(f-F)' X - s ~ v 25.4 *** 42 TABLE 2. Chi-square test of the proportions of foxtail to normal trees at Ulu Sedeli plantation Age (year) 8 Total Frequency observed, f Normal 78.2 70.4 73.0 60.9 54.0 56.0 54.0 446.5 Foxtail 11.8 19.6 17.0 29.1 36.0 34.0 36.0 183.5 Total 90.0 90.0 90.0 90.0 90.0 90.0 90.0 630.0 Frequency expected, F Normal Foxtail 63.8 26.2 63.8 26.2 63.8 26.2 63.8 26.2 63.8 26.2 63.8 26.2 63.8 26.2 (f-F)' Normal Foxtail 3.25 7.91 0.68 1.66 1.33 3.23 0.13 0.32 1.50 3.67 0.95 2.32 1.50 3.67 32.1 *** 43 TABLE 3. Chi-square test of the proportions of sub-terminal to terminal foxtails at Kemasul plantation Age (year) Total Frequency Sub-terminal 46.4 57.4 45.6 49.8 70.6 79.5 75.8 425.1 observed, f Terminal 43.6 32.6 44.4 40.2 19.4 10.5 14.2 204.9 Total 90.0 90.0 90.0 90.0 90.0 90.0 90.0 630.0 Frequency Sub-terminal 60.7 60.7 60.7 60.7 60.7 60.7 60.7 expected, F Terminal 29.3 29.3 29.2 29.3 29.3 29.3 29.3 (f-FV Sub-terminal 3.36 0.17 3.76 1.96 1.61 5.82 3.76 Terminal 6.98 0.37 7.78 4.05 3.34 12.06 7.78 X2 = 7<f-F> F 63.8 *** 44 TABLE 4. Chi-square test of the proportions of sub-terminal to terminal foxtails at Ulu Sedeli plantation Age (year) 8 Total Frequency Sub-terminal 42.8 46.0 48.0 49.1 55.3 69.2 66.4 376.8 observed, f Terminal 47.2 44.0 42.0 40.9 34.7 20.8 23.6 253.2 Total 90.0 90.0 90.0 90.0 90.0 90.0 90.0 630.0 Frequency Sub-terminal 53.8 53.8 53.8 53.8 53.8 53.8 53.8 expected, F Terminal 36.2 36.2 36.2 36.2 36.2 36.2 36.2 <f-F)' Sub-terminal 2.25 1.13 0.62 0.41 0.04 4.41 2.95 Terminal 3.34 1.68 0.93 0.61 0.06 16.55 4.39 29.4 *** 45 APPENDIX II Analysis of Variance - Diameter over-bark of normal and foxtail trees Legend * - significant. 5% level of probability ** - significant. 1% level of probability *** - significant. 0.1% level of probability ns - not significant. 46 Analysis of variance for diameter over-bark of normal and f o x t a i l trees  PLANTATION: KEMASUL TABLE 1: 4 year-old Source of v a r i a t i o n df SS MS F Between phenotypes 1 13.81 13.81 4.63* Within phenotypes 486 1451.3 2.98 Total 487 TABLE 2: 5 year-old Source of v a r i a t i o n df SS MS F Between phenotypes 1 28.79 28.79 3.12 ns Within phenotypes 754 6953.25 9.22 Total 755 TABLE 3: 6 year-old Source of v a r i a t i o n df SS MS F Between phenotypes 1 0.65 0.65 0.08 ns Within phenotypes 638 4831.96 7.57 Total 639 47 TABLE 4: 7 year-old Source of variation df SS MS F Between phenotypes 1 121.51 121.51 12.1*** Within phenotypes 414 1451.3 2.98 Total 415 TABLE 5: 8 year-old Source of variation df SS MS F Between phenotypes 1 20.98 20.98 1.54 ns Within phenotypes 852 11573.29 13.58 Total 853 48 PLANTATION: ULU SEDELI TABLE 6: 3 year-old Source of variation df SS MS F Between phenotypes 1 93.96 93.96 10.31*** Within phenotypes 509 4637.13 9.11 Total 510 TABLE 7: 4 year-old Source of variation df SS MS F Between phenotypes 1 236.9 236.9 27.54*** Within phenotypes 337 2901.48 8.60 Total 338 TABLE 8: 5 year-old Source of variation df SS MS F Between phenotypes 1 430.87 430.87 17.09*** Within phenotypes 668 16836.44 25.2 Total 669 49 TABLE 9: 6 year-old Source of variation df SS MS F Between phenotypes 1 81.59 81.59 2.35 ns Within phenotypes 300 10400.39 34.66 Total 301 TABLE 10: 7 year-old Source of variation df SS MS F Between phenotypes 1 0.83 0.83 0.02 ns Within phenotypes 557 20560.38 36.9 Total 558 TABLE 11: 8 year-old Source of variation df SS MS F Between phenotypes 1 117.16 117.16 2.48 ns Within phenotypes 814 38368.76 47.13 Total 815 50 APPENDIX III Analysis of Variance - Total height of normal and foxtail trees Legend * - significant. 5% level of probability ** - significant. 1% level of probability *** - significant. 0.1% level of probability ns - not significant. 51 Analysis of variance for total height of normal and foxtail trees  PLANTATION: KEMASUL TABLE 1: 1 year-old Source of variation df SS MS F Between phenotypes 1 6.57 6.57 4.98* Within phenotypes 1194 1576.08 1.32 Total 1195 TABLE 2: 2 year-old Source of variation df SS MS F Between phenotypes 1 16.2 16.2 25.31*** Within phenotypes 1125 720 0.64 Total 1126 TABLE 3: 4 year-old Source of variation df SS MS F Between phenotypes 1 61.48 61.48 11.11*** Within phenotypes 486 2687.58 5.53 Total 487 52 TABLE 4: 5 year-old Source of variation df SS MS F Between phenotypes 1 119.64 119.64 1.43 ns Within phenotypes 754 24399.86 38.30 Total 755 TABLE 5: 6 year-old Source of variation df SS MS F Between phenotypes 1 43.74 43.74 1.14 ns Within phenotypes 638 24441.6 38.30 Total 639 TABLE 6: 7 year-old Source of variation df SS MS F Between phenotypes 1 0.96 0.96 1.00 ns Within phenotypes 414 397.44 0.96 Total 415 53 TABLE 7: 8 year-old Source of variation df SS MS F Between phenotypes 1 68.36 68.36 0.77 ns Within phenotypes 852 74822.64 87.82 Total 853 54 PLANTATION: ULU SEDELI TABLE 8: 2 year-old Source of variation df SS MS F Between phenotypes 1 24.611 24.61 153.8*** Within phenotypes 302 50.19 0.61 Total 303 TABLE 9: 3 year-old Source of variation df SS MS F Between phenotypes 1 62.22 62.22 4.14* Within phenotypes 509 7635.0 15.0 Total 510 TABLE 10: 4 year-old Source of variation df SS MS F Between phenotypes 1 0.27 0.27 0.19 ns Within phenotypes 337 471.76 1.40 Total 338 55 TABLE 11: 5 year-old Source of variation df SS MS F Between phenotypes 1 30.32 30.32 0.51 ns Within phenotypes 668 39705.92 59.26 Total 669 TABLE 12: 6 year-old Source of variation df SS MS F Between phenotypes 1 24.66 24.66 0.44 ns Within phenotypes 300 16658.28 55.52 Total 301 TABLE 13: 7 year-old Source of variation df SS MS F Between phenotypes 1 1.2 1.2 1.24 ns Within phenotypes 557 540.29 0.97 Total 558 56 TABLE 14: 8 year-old Source of variation df SS MS F Between phenotypes 1 184.24 184.24 3.58 ns Within phenotypes 814 41896.95 51.47 Total 815 57 APPENDIX IV Analysis of Variance - Specific gravity of normal and foxtail trees Legend * - significant. 5% level of probability ** - significant. 1% level of probability *** - significant. 0.1% level of probability ns - not significant. 58 Analysis of variance for specific gravity of normal and foxtail trees TABLE 1: 10 percent height level Source of variation df SS MS F Between plantations 1 0.0454 0.0454 9.45 ** Between phenotypes/plantations 2 0.0153 0.0076 1.72 ns Between trees/phenotypes/plantations 16 0.0710 0.0044 6.28 *** Error 60 0.0413 0.0007 Total 79 TABLE 2: 30 percent height : level Source of variation df SS MS F Between plantations 1 0.0163 0.0163 1.81 ns Between phenotypes/plantations 2 0.0087 0.0039 0.40 ns Between trees/phenotypes/plantations 16 0.1533 0.0096 12.0 *** Error 60 0.0456 0.0008 Total 79 TABLE 3: 50 percent height : level Source of variation df SS MS F Between plantations 1 0.0039 0.0039 0.32 ns Between phenotypes/plantations 2 0.0188 0.0094 0.75 ns Between trees/phenotypes/plantations 16 0.2007 0.0125 10.42 *** Error 60 0.0740 0.0012 Total 79 59 TABLE 4: 70 percent height level Source of variation df SS MS F Between plantations 1 0.0003 0.0003 0.02 ns Between phenotypes/plantations 2 0.0401 0.0020 0.16 ns Between trees/phenotypes/plantations 16 0.1964 0.0123 13.66 *** Error 60 0.0573 0.0009 Total 79 TABLE 5: 90 percent height level Source of variation df SS MS F Between plantations 1 0.0076 0.0076 0.98 ns Between phenotypes/plantations 2 0.0342 0.0171 2.59 ns Between trees/phenotypes/plantations 16 0.1051 0.0066 8.25 *** Error 60 0.0509 0.0008 Total 79 

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