Journal of Vegetation Science 1: 271-276, 1990 - Population IAVS; Opulus Press Uppsala.

Printed in Sweden

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Population structure and dynamics of Juniperus excelsa in Balouchistan, Pakistan

Ahmed, Moinuddin1*, Shaukat, Syed Shahid2 & Buzdar, Abdul Hafeez3
1

Department of Botany, University of Auckland, Private Bag, Auckland, New Zealand; 2 Department of Botany, University of Karachi, Karachi, Pakistan; 3 Department of Botany, University of Balouchistan, Quetta, Pakistan;
* Tel. +64 9 737999; Fax +64 9 33429

Abstract. 60 monospecific stands of Juniperus excelsa were sampled at four locations in Balouchistan. Density, basal area and height of individuals were recorded. Soils were analysed for selected physical and chemical characteristics and the degree of disturbance due to logging and burning was noted. The density of juniper trees (> 6 cm dbh) ranged from 56 to 332 stems / ha (average 174 stems / ha). Higher densities were recorded for relatively undisturbed stands and on west facing slopes. Density of seedlings and saplings (< 6 cm dbh) was strongly correlated with tree density and tree basal area. Among the edaphic variables CaCO3 was correlated with juniper density and basal area. Diameter distributions within stands were mostly skewed and unimodal with gaps appearing in large size classes. The male to female ratio was close to 1. Cross-sections of 16 trees were used to determine age and growth rate. Number of rings in trees with 20 to 30 cm dbh ranged from 95 to 221 (x = 160 38). Diameter and age were not related. Mean annual diameter increment ranged from 6 to 16 yr / cm (x = 10 3 yr / cm). It is concluded that size class gaps and low seedling / sapling densities are the consequence of anthropogenic disturbance.
Keywords: Age; Basal area; Density; Growth rate; Size frequency distribution. Nomenclature: Stewart (1972).

tion of various tree species (including junipers) in two districts of Balouchistan. No comprehensive quantitative ecological studies have yet been published on juniper forests; hence the population structure and dynamics of Juniperus excelsa in Balouchistan have remained almost unknown. Since the time of British control local people have been allowed to use juniper forests for grazing and for collecting dry wood. With the increase in population, grazing pressure and illegal cutting, the regeneration and growth of these forests is adversely affected. Overgrazing and excessive cutting have led to increased rate of soil erosion in many areas. In many stands juniper trees are dead or dying and can be seen on almost bare rock. It is likely that the prevailing disturbances will further change the juniper forests. Therefore, it is considered urgent to obtain quantitative information on population structure and natural regeneration possibilities of the mature juniper forests in Balouchistan.

The study area Juniperus excelsa is distributed between 20 9' and 30 37' N and 67 1' and 68 3' E , as well as in some isolated dry valleys from 1200 m to 3000 m a. s. l. (Rafi 1965). The area chosen for the present investigation is situated between Kuch and Chautair, Balochistan province, on the 75 km long so-called Juniper Tract. The area includes irregular and rugged ridges with steep terrain. According to Champion, Seth & Khattak (1965), the Juniper Tract falls within the dry temperate forest region. The average annual rainfall is 269 mm with a maximum of 74 mm in July and a minimum of 3 mm in January. The hottest month is July (27.4 C), the coldest is January (7.9 C). Relative humidity ranges from 35 % in January to 60 % in September. Snow occurs between November and April with a maximum (68 cm) in February (data from Ziarat).

Introduction The population structure and dynamics of trees are of interest to ecologists, foresters and silviculturists. Few studies of Juniperus excelsa have dealt with autecological or economic aspects (Ishaque 1955; Khattak 1963; Zaman, Khan & Khan 1968; Sheikh 1985). Hussain & Rizvi (1974) examined the factors affecting the distribu-

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Ahmed, M., Shaukat, S. S. & Buzdar, A. H. Results Density and basal area Density and basal area values for trees and seedlings in the four localities are summarized in Table 1. Juniper constituted 96.6 % of the stand density. Highest stand density was recorded from Kuch while the lowest was from Ziarat. However, stands at Chautair showed the highest average density. The average juniper tree (>6 cm dbh) density was 170 stems / ha with a range of 56 - 332 stems / ha. Seedling density varied considerably both between and within sites. The average seedling density was 51 / ha (range 0 - 219 / ha). Old junipers made up 20 % of the population. The basal area of Juniperus excelsa was highly variable both within and between sites, with an average of 41. 8 m2 ha1 and a range of 8.8 to 152.2 m2 ha1 (Table 1). Basal area was significantly correlated with density (r = 0.71, P < 0.001). Variation with aspect Fig. 1 shows the distribution of juniper density and basal area on slopes with different aspects. West, southwest and north facing slopes show the highest density and basal area. The pattern of basal area is slightly different, but with the same maxima and minima. Scheffe's Multiple Contrasts Test showed that juniper density and basal area on northerly slopes (N, NW and WNW) did not differ significantly from those on southerly slopes (SW, S and SE), whereas both density and basal area were significantly higher (P < 0. 01) and P <

At each of four locations 15 stands were sampled with the Point Centered Quarter Method (Cottam & Curtis 1956). In each stand 20 points were sampled at 25 m intervals along transects in a stratified random way over at least 2 ha (Ogden & Powell 1979). Phytosociological attributes were described following MuellerDombois & Ellenberg (1974). Density figures were differentiated for seedlings and saplings (< 6 cm dbh), healthy trees (> 6 cm dbh), old trees (apparently unhealthy and with > 70 cm dbh) and damaged trees (with marks of cutting or burning). Logged junipers were counted separately. The sex of individual junipers was determined in the field. Juniper density and basal area from 60 stands on different aspects were compared using Scheffe's Multiple Contrast Statistic (Sokal & Rohlf 1981). Composite soil samples, 0 - 20 cm deep, taken from each stand, were air-dried and passed through a 2 mm sieve. Percentage of sand and maximum water holding capacity of the soils were determined following the methods in Anon. (1951). Electrical conductivity, pH, amount of organic matter and CaCO3 were estimated in accordance with Jackson (1958). Cross sections at breast height of 16 randomly selected juniper trees (20 - 30 cm dbh) from the Ziarat site were prepared and age and growth rate determined (Fritts 1976). Linear regression was calculated for tree diameter against age and growth rate, and of juniper density and basal area against edaphic and topographic factors (Sokal & Rohlf 1981).

Table 1. Summary of stand characteristics of 60 stands from four localities in Balouchistan. Jun. = Juniperus excelsa; Density figures in stems / ha; B. a. = Basal area in m2ha1; X SD = Mean Standard deviation; Values for logged Juniperus obtained separately.
Ziarat X SD Altitude m 2700 130 Density 142 59 Jun. density 140 59 Jun. B.a. 32 26 Jun. seedling 58 43 Healthy Jun. 86 35 Old Jun. 35 25 Disturbed Jun. 19 18 Logged Jun. 39 42 Male Jun. 63 26 Female Jun. 66 31 Bisexual Jun. 3 3 Range 2431 56 56 9 10 43 13 0 0 40 42 0 -2901 - 289 - 289 - 100 - 168 - 166 - 111 - 72 - 129 - 115 - 126 - 9 Babakhurwari X SD Range 2652 99 185 69 171 56 44 17 49 53 130 46 31 16 10 6 38 46 67 26 78 27 3 2 2481 96 96 23 4 9 7 1 0 34 36 0 -2801 - 309 - 306 - 88 - 172 - 242 - 58 - 19 - 194 - 124 - 124 - 7 Kuch X SD 2400 159 159 30 59 122 19 18 19 56 62 2 84 64 64 14 66 54 9 12 13 25 25 4 Range 2281 64 64 14 5 45 3 1 0 24 36 0 - 2571 - 332 - 332 - 66 - 219 - 257 - 39 - 56 - 46 - 112 - 115 - 15 Chautair X SD Range 2538 219 210 61 42 138 51 19 21 76 84 5 181 67 72 16 22 59 27 14 6 24 31 4 2301 - 2881 109 - 308 107 - 308 50 - 152 0 - 67 53 - 142 10 - 135 4 - 56 0 - 26 35 - 119 45 - 142 0 - 10

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Fig. 1. Distribution of mean density and mean basal area of Juniperus excelsa (trees > 6 cm dbh) in relation to aspect. Standard error indicated for each aspect.

Fig. 2. Overall size class frequency distribution of juniper trees (>6 cm dbh) in the study area. Each dbh class covers a 11 cm interval. Height classes (2 m intervals) include individuals >2 m. Max. = maximum individuals in a size class; Min = Minimum individuals; SD = standard deviation.

0. 05 respectively) on westerly slopes (NW, WNW, W and SW) as compared with easterly ones (ENE, SE, E and NE). Density and basal area related to edaphic and topographic factors Regressions of juniper density and basal area on edaphic and topographic factors were usually linear but not significant. Significant correlation was only found with CaCO3 content and electrical conductivity in the soil (Table 2). Population structure
0.71 0.53 0.25 0.38 0.32 0.38 *** *** ** * ** **

Table 2. Relationships between juniper density (trees > 6 cm dbh or else seedlings / saplings) and basal area and significant relationships with edaphic and topographic factors. EC = electrical conductivity of the soil. Significance levels: * = P < 0.05; ** = P < 0.01; *** = P < 0.001;
Variable Regression equation y = 3.9 + 0.27x y = 13.6 + 0.38x y = 32.7 + 0.48x y = 245.6 4.70x y = 65.9 1.50x y = 128.3 + 96.33x r Sign.

Density / Basal area Density / Seedl. dens. Basal area / Seedl. dens. CaCO3 - Density CaCO3 - Basal area EC - Density

The overall size class frequency distribution of juniper is presented in Fig. 2, while Fig. 3 shows the diameter size class frequency distribution of a few stands of high and low density respectively. The lower dbh as well as the lower height classes show the highest density, both regarding mean and maximal values (Fig. 2). The

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Table 3. Diameter (dbh cm), age and radial growth rate estimates (yr / cm Standard deviation) for 16 trees from Ziarat in order of increasing dbh.

No. 01 02 03 04 05 06 07 08 09 10 11 12 13 14 15 16

Dbh 20.0 20.2 20.9 20.9 21.0 21.7 23.6 23.8 23.9 25.3 25.4 28.3 29.1 29.6 29.9 30.5

Age 128 173 189 117 136 206 95 202 184 221 201 142 132 122 148 245

Radial growth rate 9.53 11.18 12.03 8.44 10.24 16.05 5.09 13.73 11.42 13.59 11.18 6.78 6.86 6.23 9.93 13.23 0.48 0.09 1.20 0.49 0.32 1.16 0.63 1.68 1.44 0.99 0.09 0.33 0.15 0.43 0.16 1.05

Chautair sites each had as many as 25 % old juniper trees (Table 1). At Ziarat 15 % of the junipers showed evidence of branch cutting, against only 6% in Babakhurwari. Also logging intensity varies much between sites. Natural regeneration Density of seedlings varied greatly, not only among the localities but also in different stands of the same site. High mean density values were recorded from Kuch and Ziarat while stands from Chautair showed low seedling/ sapling density. On average 51 seedlings and saplings / ha were recorded (Table 1). Seedlings occurred in the vicinity of groups of parent trees and also near dense shrubby patches. Significant correlations were found between seedling density and both tree density and basal area (r = 0.53, P < 0.001; r = 0.25, P < 0.05 respectively, Table 2). Sex distribution The relative amounts of male, female and bisexual junipers varied greatly from site to site. In some stands male plants predominated while in others female plants had a greater density. Bisexual individuals were rare, constituting about 2. 3% of the total sample (Table 1). The mean density of female plants was 74 / ha against 67 ha of male individuals. The male / female ratio did not differ significantly from 1 (Chi square = 0. 35, n. s.). Sexual reproduction and clonal growth Trees (> 6 cm dbh) of Juniperus excelsa produced large numbers of berries and seeds. However, a high

Fig. 3. Size class frequency distribution of eight juniper stands. Seedling / sapling size structure (0.5 cm to 6 cm dbh) is shown in six classes; class 1: seedlings 0.5 - 1 cm; class 2 to 6: 1 cm intervals. SD = density of seedlings and saplings, TR = density of trees. Stand numbers in circles.

tallest trees (26-28 m) were recorded from Chautair. Young stands or those at high elevations mostly contained individuals of medium height (14.1 - 20 m). The dbh distribution of individual stands (Fig. 3) shows deviations from the average picture of Fig. 2. Some sites having similar tree density show different size class structures. Generally, stands with high juniper tree density had a high seedling / sapling density as well, with the seedling and saplings represented in various size classes. Most stands show one or more gaps in the size class distribution. The Babakhurwari and Kuch sites had a higher percentage of healthy juniper individuals while Ziarat and

- Population structure and dynamics of Juniperus excelsa proportion of berries were infested by insects and contained mostly hollow seeds (Ahmed, Buzdar & Shafiq in press). Besides sexual reproduction, clonal growth was observed. Often, the lower branches become prostrate and adventitious roots are produced by buried branches. Such a root system is capable of prolonging the life of the plant when the original tree dies out. Tree age and growth rate Tree age, as measured on trees ranging in diameter from 20 to 30 cm, varied greatly in trees of similar size. The average age of the 16 trees was 160 yr. Diameter was not significantly correlated with age. The growth rate, expressed as yr / cm, varied greatly among individuals of similar size (Table 3), with values ranging from 5. 09 yr / cm to 16. 05 yr / cm, but was slow on average: 10.34 yr / cm, or just about one mm increment per year. No significant relationship was found between tree diameter and growth rate.

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Discussion Earlier density figures for the Juniper Tract (Khattak 1963; Zaman, Khan & Khan 1968) are much lower than ours: 28, 6 and 53 ind. / ha respectively. The differences may be explained by assuming that the earlier estimates were based on the entire Juniper Tract, whereas our average of 170 / ha refers to 60 mature juniper stands. The variability in juniper tree and seedling / sapling density as found in this study is predominantly associated with anthropogenic and historical factors and to a lesser extent with physiographic factors. The relatively large human population in Ziarat and consequently the greater intensity of logging, cutting and grazing have presumably reduced the juniper density in this locality. Zakaullah (1978) found that the incidence of decay was higher in trees growing on southern slopes than in trees growing on north-facing slopes. In the present study there was no significant difference between these two aspects with respect to density and basal area, but now the westerly and easterly aspects differed significantly in these parameters. This cannot be explained yet. According to Sheikh (1985), the juniper forests of Balouchistan are unstable due to lack of natural regeneration. A. R. Beg (pers. comm.) believes that Juniperus excelsa is a living fossil in the area and that its forests are deteriorating rapidly. In the present study, however, seedlings and young trees in various size classes were recorded, although they will not establish in overgrazed, eroded and open areas, because they demand an undisturbed soil and shade (Ahmed, Ahmed & Anjum 1989). The lack of seedlings and saplings in certain stands

as a likely result of overgrazing, cutting and disturbance, will probably eliminate Juniperus excelsa here. Therefore prompt action is necessary to save at least some of these stands. Despite continued disturbances in the area, the size class structure in many juniper stands is normal and a sufficient number of individuals in the smaller size classes is available. This implies adequate recruitment. Despite within-stand variation and the occurrence of size-class gaps, a balanced population structure emerges on an overall basis. Such a situation has also been reported for other tree populations (Veblen & Stewart 1982; Norton 1983; Ahmed 1984). From our studies it becomes apparent that Juniperus excelsa forests in Balouchistan are no relics and they will not deteriorate, provided preservation measures will be taken. The present study indicated that bisexual plants were rare while the male / female ratio was close to 1. Ward (1982) and Faliski (1980) found that in older populations of Juniperus communis on dry nutrient-poor sites, male plants predominated, whereas in young populations female plants did. A higher frequency of females was found in the small size classes. Therefore the marginally (though not significantly) higher density of females appears to accord with these earlier findings. On the other hand, the results are contradictory to those reported by Zaman, Khan & Khan (1968) for the Juniper Tract at large, where male trees predominated. As to the age of Juniperus excelsa, our results for 2030 cm dbh trees link up well with earlier estimations on saplings. Sheikh (1985) estimated a 2. 5 cm dbh sapling to be 50 yr, while Ahmed, Ahmed & Anjum (1989) calculated 14 - 37 yr for 2 - 3 cm dbh saplings. The relationship between tree diameter and age is often used to reconstruct the past history of forest stands (e. g. Daubenmire 1968). However, the present study showed that ages of trees of similar size vary both within and between sites. Thus, such reconstructions cannot be made without a careful check of the relation between age and size. Although Juniperus excelsa was mentioned earlier as an extremely slow growing tree (Sheikh 1985), the present study provides the first quantitative evidence for a slow growth of about one mm per year. In conclusion, the Juniperus excelsa forests of Balouchistan are neither in an unstable state, nor generally deteriorating as described earlier. Still, increased human disturbance will eventually threaten them and therefore proper management is absolutely necessary to save these forests.

Acknowledgements. We are grateful to Mr. Mohammad Rafi, Chief Conservator of the Balouchistan Forest Department for providing accommodation at Ziarat during the field trip. We

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Jackson, M. L. 1958. Soil chemical analysis. Prentice-Hall, Engelwood Cliffs, New Jersey. Khattak, G. M. 1963. Working plan for the Juniper forests of Quetta Civil Division. Govt. Printing Press, West Pakistan. Mueller-Dombois, D. & Ellenberg, H. 1974. Aims and methods of vegetation ecology. Wiley and Sons, New York. Norton, D. A. 1983. A dendroclimatic analysis of indigenous tree species, South Island, New Zealand. Ph. D. thesis, University of Canterbury, New Zealand. Ogden, J. & Powell, J. A. 1979. A quantitative description of the forest vegetation on an altitudinal gradient in Mount Field National Park, Tasmania and a discussion of its history and dynamics. Aust. J. Ecol. 4: 293-325. Rafi, M. 1965. Vegetation types of Balouchistan Province (Pakistan). Govt. Printing Press, Punjab. Sheikh, I. S. 1985. Afforestation in Juniper forests of Balouchistan. Pakistan Forest Institute, Peshawar. Sokal, R. R. & Rohlf, F. J. 1981. Biometry, 2nd ed. W. H. Freeman and Company. New York. Stewart, R. R. 1972. An annotated catalogue of the vascular plants of Pakistan. In: Nasir, E. & Ali, S. (eds.), Flora of Pakistan. Karachi University Publ., Karachi. Veblen, T. T. & Stewart, G. H. 1982. On the conifer regeneration gap in New Zealand. The dynamics of Libocedrus bidwillii stands on South Island. J. Ecol. 70: 413-436. Ward, L. K. 1982. The conservation of Juniper. Longevity and old age. J. Appl. Ecol. 19: 917-928. Zakaullah. 1978. Decay in Ziarat Juniper Forests of Balouchistan. Pak. J. For. 28: 28-34. Zaman, M. B., Khan, A. A. & Khan, M. S. 1968. Survey of Juniper berries in Balouchistan forests and prospects for their exploitation. Pak. J. For. 18: 503-509. Received 16 July 1989; Revision received 30 December 1989; Accepted 23 February 1990.

extend our thanks to Dr. John Ogden for commenting on drafts of the manuscript, and to Marion France for typing the manuscript. References Anon. 1951. Soil survey manual. U. S. Dept. of Agriculture Handbook No. 18 Washington, D. C. Ahmed, M. 1984. Ecological and dendrochronological studies on Agathis australis Salisb. (Kauri). Ph. D. thesis, University of Auckland, New Zealand. Ahmed, M., Ahmed, I. & Anjum, P. I. 1989. A study of natural regeneration of Juniperus excelsa M. Bieb. in Balouchistan, Pakistan. Pak. J. Bot. 21: 118-127. Ahmed, M. , Buzdar, A. H. & Shafiq, M. in press. Pattern of change in seed characteristics of Juniperus excelsa M. Bieb. in Balouchistan, Pakistan. Pak. J. Agri. Res. 10 (2). Champion, G. H., Seth, S. K. & Khattak, G. M. 1965. Forest types of Pakistan. Pakistan Forest Institute, Peshawar, Pakistan. Cottam, G. & Curtis, J. T. 1956. The use of distance measures in phytosociological sampling. Ecology 37: 451-461. Daubenmire, R. E. 1968. Plant communities. A text book of plant synecology. Harper and Row, New York. Faliski, J. B. 1980. Vegetation dynamics and sex structure of the population of pioneer dioecious woody plants. Vegetatio 43: 23-38. Fritts, H. C. 1976. Tree rings and climate. Academic Press, London. Hussain, Z. Z. & Rizvi, S. H. R. 1974. Ecological study of Pakistan in Quettas-Pishin and Sibi districts. Agri. Pak. 25: 165-175. Ishaque, S. M. 1955. Exploitation of Balouchistan Juniper (Juniperus macropoda) for the pencil industry. Pak. J. For. 4: 250-260.