A JOURNAL ON TAXONOMIC BOTANY, PLANT SOCIOLOGY AND ECOLOGY 12(3) REINWARDTIA A JOURNAL ON TAXONOMIC BOTANY, PLANT SOCIOLOG Y AND ECOLOG Y Vol. 12(3): 205-259. 22 Desember 2006 Editors ELIZABETH A. WIDJAJA, MIEN A. RIFAI, SOEDARSONO RISWAN, JOHANIS P. MOGEA Correspondence and subscriptions of the journal should be addressed to HERBARIUM BOGORIENSE, BIDANG BOTANI, PUSAT PENELITIAN BIOLOGI - LIP1, BOGOR, INDONESIA REINWARDTIA Vol 12, Part 3, pp: 237 – 255 RECOVERY OF A LOWLAND DIPTEROCARP FOREST TWENTY TWO YEARS AFTER SELECTIVE LOGGING AT SEKUNDUR, GUNUNG LEUSER NATIONAL PARK, NORTH SUMATRA, INDONESIA DOLLY PRIATNA* Leuser Management Unit, Jl. Dr. Mansyur No. 68 Medan 20154, North Sumatra, Indonesia KUSWATA KARTAWINATA** UNESCO Office Jakarta, Regional Science Bureau for Asia and the Pacific, Jl. Galuh (II) No. 5, Kebayoran Baru, Jakarta, P.O. Box 1273/JKT, Jakarta 12110, Indonesia. k.kartawinata@unesco.org ROCHADI ABDULHADI Herbarium Bogoriense, Pusat Penelitian Biologi, LIPI, Jl. Juanda 22, Bogor 16122, Indonesia ABSTRACT PRIATNA, D.; KARTAWINATA, K.; ABDULHADI, R. 2004. Recovery of a lowland dipterocarp forest twenty two years after selective logging at Sekundur, Gunung Leuser National Park, North Sumatra, Indonesia. Reinwardtia 12 (3): 237–255. — A permanent 2-ha plot of lowland forest selectively logged in 1978 at Sekundur, Gunung Leuser National Park, which is also a Biosphere Reserve and a World Heritage Site, North Sumatra, was established and investigated in 1982. It was re-examined in 2000, where remeasurement and reidentification of all trees with DBH ≥10 cm were made. The areas of gap, building and mature phases of the canopy were also measured and mapped. Within this plot, 133 species, 87 genera and 39 families were recorded, with the total number of trees of 1145 or density of 572.5/ha. Euphorbiaceae was the richest family with 18 species (13.5 % of the total) and total number of trees of 248 (21.7 % of the total or density of 124 trees/ha. The most important families were Dipterocarpaceae with IV (Importance Value) = 52.0, followed by Euphorbiaceae with IV = 51.8. The most prevalent species was Shorea kunstleri (Dipterocarpaceae) with IV =24.4, followed by Macaranga diepenhorstii (Euphorbiaceae) with IV = 12.4. They were the species with highest density, 34 trees/ha and 23.5 trees/ha, respectively. During the period of 18 years there has been no shift in the richest families, most important families and most important species. Euphorbiaceae was the richest family and Dipterocarpaceae was the most important family, with Shorea kunstleri as the most important species with highest importance value throughout the period. The number of species increased from 127 to 133 with increase in density by 36.8% , from 418.5 trees/ha to 572.5 trees/ha. The mortality was 25.57 % or 1.4 % per year. The diameter class distribution indicated that the forest recovery has not been complete. Trees were small, comprising 67.6 % with diameters of 10-20 cm and only two trees had diameters of 100 cm, i.e. Melanochyla caesia and Lithocarpus urceolaris. Based on the basal area of all species, the logged-over forest at Sekundur is estimated to reach the situation similar to undisturbed primary forest in 56 years after logging, but on the basis of basal area of Dipterocarpaceae such condition could be achieved in 172 years. The canopy has not fully recovered and the complete closure of gaps is estimated to take 53 years since the logging started. The canopy consisted of gap phase (24.6 %), building phase (19.7 %) and mature phase (55.7 %). During the period of 18 years the tree mortality was 25.57 % or the rate of 1.4 %/year. Euphorbiaceae experienced the highest mortality, particularly among the trees with diameters of 10-20 cm. Mortality decreased with the increase of diameters. During the same period 520 new trees of 16 species were recruited. The densities of 53 % of the species experienced changes of only one tree or no changes at all. Drastic increase in tree population occurred in light demanding species, such as Baccaurea kunstleri, Endospermum diadenum, Mallotus penangensis, Sapium baccatum and Macaranga diepenhorstii . Key words: Forest recovery, selective logging, structure and composition, mortality, recruitment, canopy closure, Sumatra. * Present Address: The Zoological Society of London, Jambi Tiger Project, , PO Box 2000, Jambi, Indonesia, dpriatna@asiaticpersada.com. ** Concurrently Research Associate at: (1) Herbarium Bogoriense, Pusat Penelitian Biologi, LIPI, Jl. Juanda 22, Bogor 16122, Indonesia, kkjak@indo.net.id; and (2) Botany Department, Field Museum, 1400 S. Lake Shore Drive, Chicago, Illinois 60605- 2496, USA 237 mailto:kkjak@indo.net.id 238 REINWARDTIA [VOL.12 ABSTRAK PRIATNA, D.; KARTAWINATA, K; ABDULHADI, R. 2004. –– Pemulihan hutan pamah dipterocarpaceae 22 tahun setelah tebang pilih di Sekundur, Taman Nasional Gunung Leuser, Sumatra Utara, Indonesia. Reinwardtia 12(3): 237–255. — Sebuah petak permanen seluas dua hektar dalam hutan pamah yang ditebang-pilih tahun 1978 dibuat dan ditelah pada tahun 1982 di Sekundur, Taman Nasional Gunung Leuser, Sumatera Utara. Petak tersebut diteliti ulang pada tahun 2000; semua pohon dengan diameter setinggi dada ≥10 cm, ditandai, diukur diameter dan tingginya dan diidentifikasi. Luas fase-fase rumpang, membangun dan matang dalam kanopi diukur dan dipetakan. Dalam petak ini tercatat 133 jenis, 87 marga dan 39 suku, dengan jumlah pohon sebanyak 1145 atau kerapatan 572.5 pohon/ha. Euphorbiaceae merupakan suku terkaya dengan 18 jenis (13.5 % dari semua jenis) dan jumlah pohon sebanyak 248 (21.7 % dari total) atau kerapatan 124 pohon/ha. Suku yang paling penting adalah Dipterocarpaceae (Nilai Penting, NP = 52.0), diikuti oleh Euphorbiaceae (NP = 51.8). Jenis yang paling menonjol adalah Shorea kunstleri (Dipterocarpaceae) dengan NP =24.4, diikuti oleh Macaranga diepenhorstii (Euphorbiaceae) dengan NP = 12.4, dan dua jenis ini mempunyai kerapatan tertinggi, masing-masing 34 pohon/ha and 23.5 pohon /ha. Selama 18 tahun tidak terdapat pergeseran suku-suku terkaya dan terpenting serta jenis-jenis terpenting. Euphorbiaceae merupakan suku terkaya dan Dipterocarpaceae suku terpenting, dengan Shorea kunstleri sebagai jenis terpenting selama 18 tahun ini. Jumlah jenis bertambah dari 127 menjadi 133 dengan peningkatan kerapatan sebanyak 36.8 %, yaitu dari 418.5 pohon/ha menjadi 572.5 pohon/ha. Mortalitas tercatat 25.57 % atau 1.4 % per tahun. Sebaran kelas diameter menunjukkan bahwa pemulihan hutan belum lengkap. Sebagian besar pohon-pohon berukuran kecil; 67.6 % termasuk kelas diameter 10-20 cm dan hanya dua pohon yang mempunyai diameter > 100 cm, yaitu Melanochyla caesia and Lithocarpus urceolaris. Berdasarkan luas bidang dasar semua jenis, hutan bekas pembalakan ini akan mencapai kondisi seperti hutan primer yang tidak terganggu dalam waktu 56 tahun setelah pembalakan, tetapi berdsarkan luas bidang dasar Dipterocarpaceae pemulihan ini memerlukan waktu 172 tahun. Kanopi hutan belum sepenuhnya pulih dan penutupan rumpang spenuhnya diperkirakan memerlukan waktu 53 tahun sejak hutan dibalak. Kanopi terdiri atas fase rumpang (24.6 %), fase membangun (19.7 %) dan fase matang (55.7 %). Selama 18 tahun mortalitas mencapai 25.57 % atau laju mortalitas 1.4 %/tahun dan tidak ada mortalitas dalam 44.1 % dari jenis. Penambahan pohon baru tercatat sebanyak 520 pohon yang termasuk16 jenis. Sebanyak 53 % dari semua jenis, kerapatannya mengalami perubahan hanya satu pohon atau sama sekali tidak mengalami perubahan. Jumlah pohon yang meningkat drastis terjadi pada jenis- jenis yang memerlukan cahaya, seperti Baccaurea kunstleri, Endospermum diadenum, Mallotus penangensis, Sapium baccatum and Macaranga diepenhorstii. Kata kunci: Pemulihan hutan, pembalakan selektif, struktur dan komposisi, mortalitas, rekrutmen, penutupan kanopi, Sumatera. INTRODUCTION In view of continuous and rapid decrease of the tropical forest area, information on forest is badly needed (Wich et al. 1999). Research on the ecology of primary forests of Indonesia is still relatively meager, although at the same time ecological information has to be accumulated before the primary forest disappears (Abdulhadi et al. 1998). During the last three decades, various ecological research activities have been undertaken by various national and international institutions, but most of them have been short- term research projects on vegetation in Kalimantan and Sumatra (Kartawinata, 1990; Lamounier, 1997). The success of conservation and management of tropical forests depends among others on a profound knowledge regarding forest dynamics (Hartshorn, 1990). To study forest dynamics, several permanent plots have been established in various localities, including at the Gunung-Gede Pangrango National Park, West Java; Kayan Mentarang National Park, Lempake and Wanariset Samboja in East Kalimantan; Gunung Palung National Park in West Kalimantan; Barito Ulu in Central Kalimantan; and the Gunung, Leuser National Park, North Sumatra (Budiman & Abdulhadi, 1995; Kartawinata 1990; Riswan, 1987). Indonesia has extensive areas of logged-over forests and degraded lands arising from intensive exploitation of forest resources. In 2000 the logged-over forests covered about 23 million hectares or 55 % of the total logging concession area (Kartawinata et al. 2001). Selective logging operations led to the formation of canopy openings, resulting from tree felling, skid trails, haul roads and log-yards. The structure and composition of residual stands have been investigated by various authors (e.g. Abdulhadi et al. 1981; Bertault et al., 1997; Cannon et al., 1994; Haeruman 1978; Rosalina 1986; Sist et al., 2003; Soemarna & Suyana 1979; Soemarno, 2001; Tinal & Palinewen 1978; see also some articles in Sist et al. 1997). The number of tree 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 239 species in logged-over forests is usually lower than in primary forests ( Kartawinata et al. 2001) and the tree mortality is higher (Cannon et al. 1994), which can be 2.1 % per year in logged over forests and 1.7 % in primary forests (Whitmore, 1984). Bare areas in the logged-over forests covered 14 to 50 % of the ground and were invaded by light-demanding, fast-growing and light-wood pioneer species (Abdulhadi et al. 1981; Fox 1969; Kartawinata et al., 1983; Meijer, 1970; Nicholson, 1958; Riswan & Kartawinata, 1991; Tinal & Palinewen 1978). Should there be no additional disturbances, logged-over forests will return to compositional and structural characteristics similar to undisturbed primary forests in at least 150 years (Riswan et al. 1986; Riswan & Kartawinata, 1988, 1991). Selective logging operations have left a mosaic of unlogged and logged areas. The unlogged primary forest areas are mainly on the less accessible or less productive areas, while the logged areas developed into secondary forest. There is a great spatial variation in term of degree of damages and consequent forest structure and species composition in the logged-over forest. This heterogeneity of habitat can support a diversity of species different from pre-logging conditions and is of value to conservation (Cannon et al. 1994). In 1982 a study of a two-hectare plot of lowland forest selectively logged in 1978 was conducted at Sekundur, Gunung Leuser National Park, North Sumatra (Abdulhadi et al., 1987). The present study was the re-invetigation of the same two-hectare plot carried out in 2000 to provide information on the recovery of selectively logged-over forests, with the objective of investigating the structural and compositional changes during the last 18 years since the previous study was conducted in 1982. STUDY AREA AND METHOD The study was carried out in a selectively logged lowland dipterocarp forest at 04 o 58’- 04 o 59’ N dan 98 o 04’-98 o 05’ E, in Sekundur, within the Gunung Leuser National Park (GLNP) and Leuser Ecosystem Area in the Besitang Sub- district, Langkat District, North Sumatra, (Figure 1). In 1981 the GLNP was designated as a Biosphere Reserve and in July 2004, together with the Kerinci Seblat and Bukit Barisan Selatan National Park, it was inscribed in the World Heritage List as the Cluster Tropical Rainforest Heritage of Sumatra. The study area is located at 75-100 m above sea level within the 1978 logging block of the PT Raja Garuda Mas (Abdulhadi et al., 1987). The terrain ranges from undulating to somewhat hilly with gentle to steep slopes. The climate is very humid without dry months with the rainfall type A (Schmidt & Ferguson, 1951), and the annual rainfall is 3500 - 4000 mm (Leuser Management Unit, unpublished). Soil in the area is classified as ‘tropudult’ (USDA) or equivalent to Red Yellow Podsolic soil (Soepraptohardjo & Ismangun 1980). The parent material is acid tuff, sandstone and sand deposit. Solum is thick, red to yellow, with variable texture, firm to friable consistency, acidic, low nutrient content, slow to medium permeability, and easily erodable . A permanent plot was subjectively established in a selectively logged-over forest in 1982 by Abdulhadi et al. (1987). The plot was two hectares (100 x 200 m) and was divided into subplots of 10 x 10 m. It covered the logging roads, skid trails, extracted area and undisturbed section of the logged forest. All trees with DBH ≥ 10 cm occurring within the subplots were recorded and numbered with metal tags at 160 cm above ground. The DBHs were measured at 130 cm above ground. For trees with tall buttresses measurements were made 20 cm above the upper ends of the buttresses. The gap, building and mature phases of the canopy (sensu Whitmore 1984) and a profile diagram were drawn. In February-March 2000 and August- November 2000 the above two-hectare permanent plot was re-surveyed and all trees were re- numbered, re-marked and re- measured. The height of each tree within the plot was measured and its position was drawn on a graph paper with the scale of 1:200. The mosaic of the gap, building and mature phases of the canopy and a profile diagram of the forest were re-drawn on a 10 x 60 m plot. Voucher specimens were collected for identification at the Herbarium Bogoriense. The density, frequency, and dominance as measured by basal area and their relative values as well as the Importance Values (Bray & Curtis, 1957) of each species were computed following the standard calculation described in detail in Mueller-Dombois and Ellenberg 1974. Calculation of the Family Importance Value follows the method used by Kartawinata et al. (2004). To show the diversity of tree species of the 1982 and 2000 results, Shannon-Winner Indices of Diversity and Evenness were calculated using the standard formulas (Magurran, 1988; Zar, 1996). 240 REINWARDTIA [VOL.12 The results of the present study were compared with the data of investigation of the same plot carried out in 1982 (unpublished data of Abdulhadi and Abdulhadi et al. 1987) and that in undisturbed primary forest at Ketambe (Abdulhadi et al. 1989). The estimate of floristic and structural recovery rate after logging was carried out by comparing the density, basal area and canopy coverage by applying the method used by Abdulhadi (1992). RESULTS AND DISCUSSION Composition In the study of the 2-ha plot in year 2000, 133 species, 87 genera and 39 families of trees with DBH ≥ 10 cm (Table 1; see Appendix 1 for details) were recorded. Eighteen years earlier in 1982, 127 species, and 88 genera, 43 families were registered (Abdulhadi et al., 1987). In the present study Euphorbiaceae was the richest family with 18 species (13.5% of the total), followed by Lauraceae with 10 species (7.5%), and Anacardiaceae and Dipterocarpaceae with 8 species (6%) each. Beside the richest in species, Euphorbiaceae had the highest number of genera ( 11) and number of trees (248). It is well known that, next to Dipterocarpaceae, Euphorbiaceae is in general the richest family in the primary and secondary lowland rain forests of Malesia (Abdulhadi et al. 1991; Kartawinata et al. 1981; Kartawinata, et al. 2004;Riswan, 1987). The success of Euphorbiaceae appeared to be closely related to its adaptive capability. It contains species preferring to grow on open places, such as gaps and they form the canopy of secondary forests (Riswan, 1982; Whitmore, 1984; Riswan, 1987; Manullang, 1998). Figure 1. Map of the study area in a selectively logged lowland forest at Sekundur, GLNP, North Sumatera 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 241 The Importance Values of families differed from density, frequency and dominance (basal area). Seven families had FIV (Family Importance Values) >10, where Dipterocarpaceae had the highest FIV (52.0), followed by Euphorbiaceae with FIV of 51.8. These high values were contributed by Shorea kunstleri (Diptero-carpaceae) with IV (Importance Value) of 24.4 and Macaranga diepenhorstii (Euphorbiaceae) with IV of 12.4. Table 1. Important families in the 2-ha plot of a selectively logged lowland forest at Sekundur in the Gunung Leuser National Park, North Sumatra in 1982 and 2000. Family Number of Species % Number of species Number of Genera Number of trees (DBH ≥ 10 cm) Family Importance Value 1982 * 2000 1982 * 2000 1982 * 2000 1982 * 2000 1982 * 2000 Dipterocarpaceae 7 8 5.52 6.02 3 3 118 127 57.6 52.0 Euphorbiaceae 19 18 14.96 13.53 11 11 94 248 27.5 51.8 Lauraceae 10 10 7.87 7.52 5 5 71 96 21.1 21.0 Anacardiaceae 8 8 6.29 6.02 5 5 55 65 24.2 20.0 Myrtaceae 6 6 4.72 4.51 2 2 54 64 17.8 15.4 Sapotaceae 4 4 3.15 3.01 3 3 31 59 9.2 13.0 Flacourtiaceae 4 4 3.15 3.01 4 4 41 45 13.0 11.6 Annonaceae 7 4 5.52 3.01 6 4 33 38 12.5 10.0 Fagaceae 3 3 2.36 2.26 2 2 14 21 9.4 8.9 Tiliaceae 1 1 0.79 0.75 1 1 42 40 11.8 8.6 Moraceae 3 3 2.36 2.26 2 2 29 33 9.7 8.39 Other families 55 64 43.31 48.10 44 45 255 309 86.2 87.7 Total 127 133 100 100 88 87 837 1145 300 300 *) Abdulhadi (unpublished data) Ten most important species arranged in descending order of Importance Values (Figure 2) were Shorea kunstleri, Mangifera gracilipes, Cinnamomum iners, Eugenia acutangula, Pentace polyantha, Cleistanthus bakonensis, Shorea pauciflora, Lophopetalum javanicum, Lithocarpus urceolaris and Mezzettia parviflora. The total IVs of these species was 35.5 % of the total IVs of all species and Shorea kunstleri had the highest IV of 24.4 or 8.11 % of the total. Data recorded in 1982 (Abdulhadi et al. 1987) showed similar values, where the IVs of ten most important species was 38.8 %, and S. kunstleri had the highest IV of 29.8 or 9.9 % of the total; only one species, Cleistanthus bakonensis, was secondary species in this group. It should be noted, however, that Macaranga diepenhorstii with IV of 12.37, Endospermum diadenum with IV of 7.62 and Sapium bacccatum with IV of 7.36 should be considered important species in year 2000 whereas they had lower IVs in 1982. Figure 2 also shows the decrease of the IVs in 8 of the 10 most important species when IVs in 0 5 10 15 20 25 30 35 Sk Mg Ci Ea Pp Cb Mp Sp Lj Lu Species Im p o rt a n ce V a lu e 1982 2000 Fig. 2. Ten most important species recorded in 1982 (Abdulhadi, unpublished data) and 2000 in a 2- ha plot of a selectively logged lowland forest at Sekundur, GLNP, North Sumatra. Sk= Shorea kunstleri; Mg= Mangifera gracilipes; Ci=Cinnamomum iners; Ea= Eugenia acutangula; Pp=Pentace polyantha; Cb=Cleistanthus bakonensis; Mp= Mezzettia parviflora; Sp= Shorea pauciflora; Lj= Lophopetalum javanicum; Lu= Lithocarpus urceolaris. 1982 compared with those in 2000. The decrease were likely attributed to the death of trees, where, except for Lophopetalum javanicum, the mortality rates of these species were 4.76 –51.85 %. Meanwhile, the IVs of Mezzettia parviflora and Lophopetalum javanicum increased, which could be attributed to the high percentage of recruitment of 57.1 % for the former and 44.4 % for the latter. 242 REINWARDTIA [VOL.12 Twenty two years after logging, the ten most important tree species based on density in the two-hectare permanent plots are shown in Table 2. The density of these ten tree species constituted 35.7 % of the total density of all tree species. Note that in 2000, a secondary forest species, Macaranga diepenhorstii, was the leading species with the highest density of 34 trees/ha and represented 5.9 % of the total density. The second prevailing species was Shorea kunstleri with density of 23.5 trees/ha and made up of 4.1 % of the total. It was noted that Macaranga diepenhorstii filled up and grew in the gaps created by selective logging whilst Shorea kunstleri occupied the unlogged portion of the forest within the plot. Density measurement undertaken in the same plot in 1982 (Abdulhadi, unpublished data) showed a similar pattern where the ten most important species made up 37.4 % of the total. During the period of 18 years, a drastic increase of density occurred in Macaranga diepenhorsti from 10 trees/ha in 1982 to 34 trees/ha in 2000, Sapium baccatum from 1 trees/ha to 19 trees/ha and Litsea noronhae from 0 to 15 trees/ha. Litsea noronhae should be registered as a new arrival. There was, however, a decrease in density of Shorea kunstleri during the period of 18 years. In 1982, Shorea kunstleri was the leading species with the highest density of 25.5 tree/ha or 12.2 % of the total, whereas in 2000 its density was 23.5 tree/ha. See also Table 4 for density figures of all species. Table 2 . Ten most important tree species based on density (trees/ha) in 2000 compared with the density in 1982 in the two-hectare plot of a lowland dipterocarp forest at Sekundur, Gunung Leuser National Park, North Sumatra. Density (tree/ha) Species Family Year 2000 Year 1982 * Macaranga diepenhorstii Euphorbiaceae 34 10 Shorea kunstleri Dipterocarpaceae 23.5 25.5 Eugenia acutangulum Myrtaceae 22.5 19 Cinnamomum iners Lauraceae 21 21 Pentace polyantha Tiliaceae 20 21 Sapium baccatum Euphorbiaceae 19 1 Mangifera gracilipes Anacardiaceae 17 15 Mezzettia parviflora Annonaceae 16.5 10.5 Endospermum diadenum Euphorbiaceae 16 3.5 Litsea noronhae Lauraceae 15 0 *) Abdulhadi (Unpublished data) Table 3. The ten most important tree species based on basal area ( m 2 /ha) measured in 2000 and in 1982 in the two- hectare plot of a lowland dipterocarp forest at Sekundur, Gunung Leuser National Park, North Sumatra. Basal Area (m 2 /ha) Species Family Year 1982* Year 2000 Increase in 18 years Shorea kunstleri Dipterocarpaceae 3.89 4.48 0.59 Dipterocarpus grandiflorus Dipterocarpaceae 0.91 1.25 0.34 Mangifera gracilipes Anacardiaceae 1.46 1.21 (-0.25) Lithocarpus urceolaris Fagaceae. 1.1 1.16 0.06 Shorea pauciflora Dipterocarpaceae 0.67 0.91 0.24 Macaranga diepenhorstii Euphorbiaceae 0.1 0.84 0.74 Lophopetalum javanicum Celastraceae 0.57 0.82 0.25 Shorea leprosula Dipterocarpaceae 0.46 0.79 0.33 Shorea multiflora Dipterocarpaceae 0.46 0.78 0.32 Mezzettia parviflora Annonaceae 0.53 0.74 0.21 Total 10.15 12.98 2.83 *) Abdulhadi (Unpublished data) In term of basal area the ten most important tree species measured in 2000 (22 years after selective logging) in the two-hectare plot are presented in Table 3. The total basal area of the ten most important species amounted to 12.98 m 2 /ha or 43.5 % of the total basal area of all species. Shorea kunstleri was the leading species 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 243 with the basal area of 4.48 m 2 /ha or 16.1 % of the total basal area for all species. The next important species was Dipterocarpus grandiflorus with the basal area of 1.25 m 2 /ha or 4.5 % of the total for all species. The measurements in the same plot made in 1982 (Abdulhadi, unpublished data) showed comparable figures, where the total basal area of the ten most important species amounted to 10.15 m 2 /ha or 47.2 % of the total for all species. Within the period of 18 years, with the exception of Mangifera gracilipes, the basal area increased ranging from 0.06 to 0.74 m 2 /ha. Being a fast-growing secondary forest species, Mangifera gracilipes showed the highest increase (0.74 m 2 /ha). The second highest increase was shown by Shorea kunstleri with the increase of 0.59 m 2 /ha; it may be considered as a fast growing species, faster than Shorea leprosula, which is generally accepted as one of the fast growing dipterocarps. The basal area increase of the other dipterocarp species ranged from 0.24 to 0.34 m 2 /ha. Structure The diameter class distribution of 837 trees recorded in 1982 and 1145 trees measured in 2000 (Figure 3) shows a shape almost like a short. inverted J, the shape of the curve typical for primary forest (Whitmore, 1984; Abdulhadi et al. 1991). It implies also that new growth is booming along just fine. In general the trees were small, consisting of 67.6 % with diameters of 10- 20 cm and 17.0 % of diameters 21-30 cm. Only two trees (0.2 %) had diameters > 100 cm, i.e., Melanochyla caesia (Anacardiaceae) and Lithocarpus urceolaris (Fagaceae). It can be definitely inferred that the recovery process of the selectively logged forest here is still in progress The forest has not reached the conditions of an undisturbed primary forest 22 years after logging. Within this 2-ha plot, Euphorbiaceae and Dipterocarpaceae had the highest number of trees, 248 (density = 124/ha) and 127 ( density = 63.5 trees/ha) and were greater than in 1982 (Table 1). More than 85 % of the trees of Euphorbiaceae consisted of Macaranga diepenhorstii, Sapium baccatum, Endospermum diadenum, Cleistanthus bakonensis, Mallotus penangensis and Baccaurea lanceolata, which were pioneer species filling up gaps or growing on forest edges. In Dipterocarpaceae 73 % of the trees were composed of Shorea kunstleri, S. multiflora and S. leprosula., which are primary species that usually grow better in small gaps than in open sites or under closed canopy (Abdulhadi et al. 1987). 0 100 200 300 400 500 600 700 800 900 I II III IV V VI Diameter Class N u m b e r o f T re e s Number of Trees in 1982 Number of Trees in 2000 Figure 3. Diameter class distribution of trees with DBH ≥ 10 cm recorded in 1982 (Abdulhadi, unpublished data) and 2000 in a 2-ha plot of selectively logged forest at Sekundur, GLNP, North Sumatra. I = 10-20 cm; II = 21-30 cm; III = 31-40 cm; IV = 41-50 cm; V = 51-60 cm; VI = > 60 cm In year 2000, 1145 trees with DBH ≥ 10 cm were recorded in the 2-ha plot, an ingrowth of 308 trees in 18 years. Forty species (31 % of the total) were each represented by a single tree, while 62 species (23 %) were each represented by 11 trees. Macaranga diepenhorstii and Shorea kunstleri were the most abundant species represented by 68 and 47 trees, respectively. S. kunstleri had the mean diameter greater than that of M. diepenhorstii. In the forest of Sekundur M. diepenhorstii regenerated well, especially in gaps and other open places Table 3 shows that the total basal area of 1145 trees (572.5 tree/ha) in the plot was 55.36 m 2 (27.68 m 2 /ha). Of these the Dipterocarpaceae contributed 127 trees with basal area of 16.49 m 2 (8.25 m 2 /ha) In 1982, the same plot contained 837 trees with the total basal area of 42.87 m 2 (21.44 m 2 /ha), including 118 trees of Dipterocarpaceae with basal area of 12.98 m 2 (6.49 m 2 /ha). Table 3 shows also that there was an increase of basal area from 21.44 m 2 /ha in 1982 to 27.68 m 2 /ha 18 years later in 2000, implying that the rate of basal area increment was 0.35 m 2 /year. Using this rate of increment and the total basal area of undisturbed primary forest at Ketambe of 40.90 m 2 /ha (Abdulhadi et al. 1989), it can be estimated that the logged- over forest at Sekundur would reach the structure similar to the undisturbed primary forest in 244 REINWARDTIA [VOL.12 (40,90-27.68)/0.35 = 37.77 years from year 2000 or 33.77 + 18 = 55.77 years from 1982. If the basal area of Dipterocarpaceae was used as the basis of calculation it would take 154.39 years from the year 2000 or 154.39 + 18 = 172.39 years from the year 1982. This is based on the fact that in 18 years the basal area of Dipterocarpaceae in the logged-over forest increased from 6.49 m 2 /ha in 1982 to 8.25 m 2 /ha in 2000, thus giving the rate of basal area Figure 4. The gap, building and mature phases of the canopy of a 2-ha plot of selectively logged forest 22 years after logging at Sekundur, GNLP, North Sumatra. The mature phase consists of the unlogged forest left during the logging in 1978 and the mature phase developed from the building phase during 18 years since the observation made in 1982. Figure 5. Gap, building and mature phases in the canopy of 2-ha logged-over lowland forest four years after logging at Sekundur, GLNP, North Sumatra (After Abdulhadi et al.1987). The mature phase consists of the unlogged forest left during the selective logging in 1978. 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 245 increment of 0.098 m2/year, while the basal area of Dipterocarpaceae in the undisturbed primary forest at Ketambe was 23.38 m 2 /ha (Abdulhadi et al. 1989). Meanwhile the restoration of a selectively logged forest (Meijer 1970) and a small area of clear-cut forest (Riswan et al. 1986) to a forest similar to original undisturbed conditions would take more than 150 years. However, due to various habitat changes during the log extraction, such as loss of nutrients and soil soil compaction, the logged-over forest will probably never return to original conditions. Gaps Figure 4 shows the results of mapping the canopy in the two-ha plot carried out in the year 2000 or 22 years after selective logging, indicating the gap, building and mature phases . It should be noted that the mature phase consists of the unlogged forest left during the selective logging in 1978 and the mature phase developed from the building phase during 18 years since the observation made in 1982. It is evident that there were many small and big gaps forming scattered patches with a total area of 4920 m 2 (24.6%), while the building phase and mature phase covered 3940 m 2 (19.7%) and 11140 m 2 (55.7%), respectively (Table 4). Figure 5 shows the gap, building and mature phases of the canopy in 1982 with their areas shown in Table 3. The gap area amounted to 31 %, indicating the severe damage of the canopy that affected the further development of the forest. Comparing the above canopy situations revealed that as yet 22 years after logging the full recovery has not been achieved. Gaps in undisturbed lowland primary forests of Malesia are only 10 –17 % of the canopy coverage (Hopkins et al., 1976; Partomihardjo et al., 1987; Poore, 1968; Whitmore, 1984) Table 4. Basal area of trees with DBH ≥ 10 cm in a 2-ha plot of a selectively logged lowland forest four years (1982) and 22 years (2000) after logging at Sekundur and in an undisturbed lowland primary forest at Ketambe, GLNP, North Sumatra. Four years after logging (1982)* Twenty two years after logging (2000) Undisturbed primary forest**) Number of trees/ha 418.5 572.5 538 Basal area (m 2 /ha) 21.44 27.68 40.90 Number of trees of Dipterocarpaceae/ha 59 63.5 139 Basal area of Dipterocarpaceae (m 2 /ha) 6.49 8.25 23.38 Source: *) Abdulhadi (unpublished data of 2 ha logged forest at Sekundur); **) Abdulhadi et al. (1989) from primary forest plot of 1.6 ha at Ketambe, TNGL, North Sumatra Table 5. The area and percentage of the canopy phases in a 2-ha plot of a selectively logged lowland forest four years and 22 years after logging at Sekundur, GLNP, compared with an undisturbed lowland dipterocarp forest at Sungei Menyala, Peninsular Malaysia Canopy phase Four years after selective logging (1982)* Twenty two years after selective logging (2000) Primary forest ** Area (m 2 ) % Area Area (m 2 ) % Area Area (m 2 ) % Area Gap 6200 31.0 4920 24.6 2400 12.0 Building 6200 31.0 3940 19.7 6800 34.0 Mature 7600 38.0 11140 55.7 10800 54.0 *) The present 2-ha plot measured in 1982 (Abdulhadi et al., 1987). **) A primary lowland dipteroccarp forest at Sungei Menyala (Whitmore, 1984). Table 5 shows also the closure of 20.6 % of gaps from 6200 m 2 to 4920 m 2 during the period of 18 years or a rate of closure of 1.14 % per year, while the mature phase increased by 46.6 % or a rate of 2.6 % per year. Trees that played a role in 246 REINWARDTIA [VOL.12 the closure of gaps were 34 % Euphorbiaceae (in particular Baccaurea kunstleri , Cleistanthus bakonensis, Endospermum diadenum, Macaranga diepenhorstii, Mallotus penangensis and Sapium baccatum), 9.5% Dipterocarpaceae ( especially Shorea kunstleri, S. pauciflora and S. multiflora), 8.4 % Lauraceae (particularly Cinnamomum inners and Litsea noronhae), and 5 % Anacardiaceae (in particular Mangifera gracilipes and Mangifera odorata). Other species contributing to the gap closure included Lophopetalum javanicum (Celastraceae), Archidendron bubalinum (Fabaceae), Artocarpus kemando (Moraceae), Ardisia lanceolata (Myrsinaceae), Eugenia acutangula (Myrtaceae), Pentace polyantha (Tiliaceae), and Teijsmanniodendron coriaceum (Verbenaceae). Considering the closure of gaps took place in 18 years from 6200 m 2 to 4920 m 2 or 71.11 m 2 per year and referring to the area of gaps of 2400 m 2 in an undisturbed forest of similar kind, it is predicted that the gap pattern in the logged- over forest at Sekundur would be restored to a condition similar to undisturbed forest in about 53 years after logging. The gap phase of 6200 m 2 measured in 1982 had developed into building and mature phases of 4240 m 2 (68.4% of the total gap area in 1982), while the remaining 1960 m 2 (31.6%) by 2000 still remained in gaps whose ground surfaces were invaded luxuriantly by a creeping fern Dicranopteris linearis of 1-2 m thick. The largest area of such gaps occurred on the logging roads where the D. linearis cover was gradually thinning out as the tree crowns were getting wider. While in general the total area of gaps decreased during the period of 18 years as indicated in Table 5, it was observed also that during the same period new gaps, resulted from broken crown and naturally fallen trees, were also formed totaling 2960 m 2 or 21.5 % giving the rate of formation of 1.2 % per year. This is slightly higher than 1.05 % recorded for East Kalimantan forest (Partomihardjo et al. 1987). The gap formation could be attributed to the Bohorok windstorm that regularly passed through the area. Figure 4 shows one large and several small gaps that did not develop into building phase during the period of 18 years, totaling 1960 m 2 or 31.6% of the total area in1982. They were mainly logging roads and skidtrails with bare and compacted soils devoid of top layers. Figures 6 show the profile diagrams of the logged-forest 22 years after logging. It is evident that the second layer with height of 10-20 m was already well occupied by young trees. It was apparently attributed to the growth of both undamaged and damaged trees and re-sprouting Figure 6. Profile diagram of a selectively logged lowland forest at Sekundur, GLNP, North Sumatra 22 years after logging. Shaded trees are species of Dipterocarpaceae, including Diterocarpus grandiflorus and Shorea kunstleri as the emergent reaching the height of about 40 m. 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 247 was responsible for the growth of damaged trees. Soemarno (2001) found that in Sekundur forest recovery on logging roads were slower than on skidtrails, which in turn slower than on the areas of log extraction. In Sabah, Meijer (1970) noted that such areas were still discernible 40 years after logging. The slow recovery was perhaps due to heavy disturbance of soils, whose magnitude depend on the nature of soils, topography, logging intensity, technique of logging and the size and numbers of the equipment used (Kartawinata et al. 2001). The disappearance of top soils resulted in the loss of seed bank in the soil (Abdulhadi et al., 1987). On compacted logging tracks the water infiltration rate is slow and could be seven times slower than that in the undisturbed soils (Abdulhadi et al. 1981), leading to an increase in surface runoff and subsequent erosion (Burgess 1971, Liew 1974). Growth of dipterocarp seedlings are hampered by drainage impediment resulted from soil compaction (Kartawinata et al., 2001) Changes in tree density and species richness Between 1982 and 2000, the number of trees and the species richness increased (Table 6). In 1982 there were 837 trees recorded in 2-ha plot of which 214 trees could not be recovered in 2000. It indicates that during the period of 18 years the tree mortality was 25.57% or 1.4% per year, with the highest mortality occurred in the 10-20 cm diameter class, where 118 trees (14.10 %) died (Figure 7) . It was observed also that the mortality decreased as the diameter increased. Most of the 214 trees died between 1982 and 2000, were Euphorbiaceae (16.8 %), including Cleistanthus bakonensis and Macaranga diepen- horstii, followed by Tiliaceae (8.9 %), i.e., Pentace polyantha, while Anacardiaceae (M. odorata) and Dipterocarpaceae (Shorea kunstleri) lost only 8.9 %, respectively. The mortality rate in the present study area was lower than that of the result of a long term investigation in undisturbed forest at Ketambe, which was only 2.3% per year (Wich et al. 1999). The high mortality at Ketambe was attributed among others to the high density of the strangling figs (Schaik, 1996), which was 8.5 trees/ha (Palombit, 1992). The mortality at Sekundur was comparable to the rate of 1-2 % generally recorded in tropical forests elsewhere (Swaine et al., 1987; Whitmore, 1984), although lower than the rate of 2.1 % per year occurring in secondary forests, where 40 % of the mortality taking place in trees with DBH of 19-24 cm (Whitmore, 1984). 0 20 40 60 80 100 120 140 10--20 20--30 30--40 40--50 50--60 60--70 70--80 80--90 90--100 > 100 Diameter class (cm) M o rt a lit y Figure 7. Mortality of trees by diameter classes between 1982 and 2000 in the 2-ha plot of selectively logged lowland forest at Sekundur, GLNP, North Sumatra. Figure 8 shows that there was no mortality in 56 of 127 species recorded in 1982 (Class I). The remaining 71 species experienced mortality Table 6. Changes in composition and density of trees with DBH ≥ 10 cm in the 2-ha plot of a selectively logged lowland forest between 1982 (Abdulhadi, unpublished) and 2000 at Sekundur, GLNP, North Sumatra. Four years after logging (1982) Twenty two years after logging (2000) Number of trees 837 1145 Number of species 127 133 Number of genera 88 87 Number of families 40 39 Species diversity index (H’) 1.826 1.843 Species evenness index (E) 0.271 0.262 248 REINWARDTIA [VOL.12 between 10 % and 100 % . Sixteen species had 100 % mortality (Class VII). Ten of them did not regenerate. Single trees that did not regenerate included Alstonia sp. (Apocynaceae), Dillenia indica (Dilleniaceae), Macaranga triloba, Spathiostemon javensis (Euphorbiaceae), Petunga sp. (Rubiaceae), Polyalthia sumatrana, Popowia hirta, Xylopia mucronata (Annonaceae), Scaphium macropodum (Sterculiaceae), and Scleropyrum cf. wallichianum (Santalaceae). 0 10 20 30 40 50 60 I II III IV V VI VII Mortality class N u m b e r o f sp e ci e s Fig. 8. Number of species and mortality class (I= 0.0%; II= 1.0-19.9 %; III= 20.0 -39.9 %; IV= 40.0-59.9 %; V= 60.0-79.9 %; VI= 80.0-99.9 %; VII= 100%) in the 2-ha plot of selectively logged lowland forest during the period of 18 years at Sekundur, GLNP, North Sumatra. In 18 years, 520 new trees with DBH of 10-46 cm belonging to 101 species appeared in the 2-ha plot of the selectively logged forest. Among these new recruits, 16 species with a total of 17 trees were not recorded in 1982, indicating new appearance stimulated by logging. The 16 new species included the following: Barringtonia macrostachya (Lecythidaceae), Canarium kipella (Burseraceae), Dysoxylum sp3., Dysoxylum sp4. (Meliaceae), Elattostachys sp. (Sapindaceae), Euodia robusta, Euodia sp1. (Rutaceae), Garcinia dioica (Clusiaceae), Leea sp. (Leeaceae), Myristica maxima (Myristicaceae), Rubiaceae sp1. (Rubia- ceae), Shorea sp2. (Dipterocarpaceae), Sizygium racemosum (Myrtaceae), Trigonostemon serratus (Euphorbiaceae), Vitex gamosepala (Verbenaceae) and Xanthophyllum erhychum (Polygalaceae). The number of species in secondary forests within the selectively logged forests is less than in primary forests. Figure 9 shows the changes of number of trees in 127 species during the period of 18 years. It should be noted that 52.8 % of the species showed the change in density only by one tree or no change at all. The number of trees of eight species (Ganua mottleyana , Mezzettia parviflora, Litsea noronhae, Baccaurea kunstleri, Mallotus penangensis, Endospermum diadenum, Sapium baccatum and Macaranga diepenhorstii) changed from 12 to 48 trees. The number of trees of light- demanding species (Baccaurea kunstleri, Mallotus penangensis Endospermum diadenum, Sapium baccatum and Macaranga diepenhorstii) in- creased sharply. Most likely they were recruited from seeds stored in the soils under the canopy in response to the formation of gaps in the canopy. Figure 9. The change in number of trees during the 18 year period in 127 species occurring in a 2-ha plot of the selectively logged lowland forest at Sekundur, GLNP, North Sumatra. 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 249 CONCLUSION During the period of 18 years there has been no shift in the richest families, most important families and most important species. Euphorbiaceae was the richest family and Dipterocarpaceae was the most important family. Shorea kunstleri was the most important species with highest importance values throughout the period. The number of species increased from 127 to 133 with increase in density by 36.8%. Euphorbiaceae experienced the highest mortality, particularly among the trees with smaller diameters. Mortality decreased with the increase of diameters. In 18 years the tree mortality rate was 1.4 % per year. The diameter class distribution indicated that the forest recovery has not been fully achieved. The canopy has not fully recovered and the complete closure of gaps is estimated to take 58 years since the logging started. Based on the basal area of all species, the logged-over forest at Sekundur is estimated to reach a situation similar to undisturbed primary forest in 56 years after logging, but on the basis of basal area of Dipterocarpaceae such condition could be achieved within 172 years. The construction of wide logging roads and skidtrails and heavy compaction of soils delayed the recovery of the logged-over forest. The above facts have implications for the improvement of silvicultural system by adopting the reduced-impact logging technique in order to reduce the degree of destruction, hence increase the recovery rate and thus reduce the length of cutting cycle. Without additional disturbances the selectively logged forest will naturally develop into a more complex forest through succession. The recovery may be accelerated by rehabilitation measures while allowing natural succession to take place. In the Sekundur forest the objective of rehabilitation should be to achieve species diversity, hence the use of a wider set of species should be preferred. ACKNOWLEDGEMENT We are grateful to all persons, including the reviewers, who assisted us in many ways that made this study and publication possible. We thank the Head of the Gunung Leuser National Park for the permit to undertake this study in the Park. The first author (DP) received a financial assistance and other supports from the Leuser Development Program, for which he wishes to render his thanks to the Director and staff of the Leuser Management Unit, in particular Prof. M. Ali Basyah Amin, Mr. Mike Griffiths, Dr. Zainal A. Pian, Dr. Kathryn A. Monk, and Dr. Yarrow Robertson. 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Reduced-impact logging in Indonesian Borneo: some results confirming the need for new silvicultural prescriptions. Forest Ecology and Management 179: 415-427. ZAR, J.H. 1996. Biostatistical analysis. Prentice-Hall International Inc., New Jersey. 252 REINWARDTIA [VOL.12 Appendix 1. Composition of tree species with DBH ≥ 10 cm in a two-hectare plot of selectively logged lowland forest at Sekundur, Gunung Leuser National Park, North Sumatra. Family and Species Basal area (m²) Density (trees/ha) Frequency (%) Relative Density (%) Relative Frequency (%) Relative Basal Area (%) Importance Value (%) 1 2 3 4 5 6 7 8 1. Anacardiaceae: 4.70 32.50 0.29 5.677 5.734 8.487 19.899 1 Campnospermum auriculatum 0.09 2.50 0.020 0.437 0.402 0.164 1.003 2 Dracontomelon dao 0.07 0.50 0.005 0.087 0.101 0.127 0.315 3 Gluta renghas 0.10 4.00 0.040 0.699 0.805 0.172 1.676 4 Mangifera foetida 0.04 1.50 0.015 0.262 0.302 0.070 0.634 5 Mangifera gracilipes 2.41 17.00 0.140 2.969 2.817 4.354 10.141 6 Mangifera odorata 0.42 4.50 0.040 0.786 0.805 0.764 2.355 7 Mangifera sp1. 0.14 2.00 0.020 0.349 0.402 0.250 1.001 8 Melanochyla caesia 1.43 0.50 0.005 0.087 0.101 2.585 2.773 2. Annonaceae: 1.67 19.00 0.18 3.319 3.622 3.017 9.958 9 Cananga odorata 0.05 1.00 0.010 0.175 0.201 0.088 0.464 10 Goniothalamus giganteus 0.01 0.50 0.005 0.087 0.101 0.021 0.209 11 Mezzettia parviflora 1.47 16.50 0.155 2.882 3.119 2.655 8.655 12 Polyalthia lateriflora 0.14 1.00 0.010 0.175 0.201 0.254 0.630 3. Apocynaceae: 0.33 1.50 0.015 0.262 0.302 0.591 1.155 13 Dyera costulata 0.33 1.5 0.015 0.262 0.302 0.591 1.155 4. Bombacaceae: 0.71 12.50 0.120 2.183 2.414 1.284 5.882 14 Durio griffithii 0.71 12.50 0.120 2.183 2.414 1.284 5.882 5. Burseraceae: 0.66 9.00 0.090 1.572 1.811 1.201 4.583 15 Canarium caudatum 0.56 5.00 0.050 0.873 1.006 1.007 2.886 16 Canarium kipella 0.02 0.50 0.005 0.087 0.101 0.028 0.216 17 Dacryodes laxa 0.03 0.50 0.005 0.087 0.101 0.053 0.241 18 Dacryodes rostrata 0.01 1.00 0.010 0.175 0.201 0.019 0.395 19 Santiria oblongifolia 0.05 2.00 0.020 0.349 0.402 0.094 0.846 6. Celastraceae: 1.64 13.00 0.125 2.271 2.515 2.959 7.745 20 Lophopetalum javanicum 1.64 13.00 0.125 2.271 2.515 2.959 7.745 7. Clusiaceae: 0.87 8.50 0.085 1.485 1.710 1.571 4.766 21 Calophyllum saigonense 0.38 2.00 0.020 0.349 0.402 0.680 1.432 22 Calophyllum soulattri 0.29 1.00 0.010 0.175 0.201 0.518 0.894 23 Calophyllum venulosum 0.11 3.50 0.035 0.611 0.704 0.194 1.510 24 Garcinia celebica 0.02 0.50 0.005 0.087 0.101 0.040 0.228 25 Garcinia dioica 0.01 0.50 0.005 0.087 0.101 0.014 0.202 26 Garcinia havilandii 0.07 1.00 0.010 0.175 0.201 0.125 0.501 8. Dipterocarpaceae: 16.49 63.50 0.550 11.092 11.066 29.790 51.948 27 Dipterocarpus grandiflorus 2.49 5.50 0.055 0.961 1.107 4.496 6.564 28 Dipterocarpus rigidus 0.07 1.50 0.015 0.262 0.302 0.118 0.682 29 Hopea beccariana 0.01 0.50 0.005 0.087 0.101 0.024 0.212 30 Shorea kunstleri 8.96 23.50 0.205 4.105 4.125 16.184 24.414 31 Shorea leprosula 1.57 10.50 0.090 1.834 1.811 2.843 6.488 32 Shorea multiflora 1.55 12.50 0.090 2.183 1.811 2.808 6.802 33 Shorea pauciflora 1.82 9.00 0.085 1.572 1.710 3.287 6.570 34 Shorea sp2. 0.02 0.50 0.005 0.087 0.101 0.030 0.218 9. Ebenaceae: 0.30 9.00 0.090 1.572 1.811 0.550 3.933 35 Diospyros malabarica 0.15 3.00 0.030 0.524 0.604 0.276 1.404 36 Diospyros pychocarpa 0.15 6.00 0.060 1.048 1.207 0.274 2.529 10. Euphorbiaceae: 6.84 124.00 0.885 21.659 17.807 12.347 51.813 37 Aporusa antennifera 0.03 1.00 0.010 0.175 0.201 0.058 0.434 38 Aporusa nitida 0.18 2.50 0.025 0.437 0.503 0.334 1.274 39 Aporusa quadrilocularis 0.03 1.50 0.015 0.262 0.302 0.049 0.613 40 Baccaurea deflexa 0.24 3.50 0.030 0.611 0.604 0.431 1.646 41 Baccaurea kunstleri 0.71 11.00 0.105 1.921 2.113 1.286 5.320 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 253 Appendix 1. continued. Family and Species Basal area (m²) Density (trees/ha) Frequency (%) Relative Density (%) Relative Frequency (%) Relative Basal Area (%) Importance Value (%) 1 2 3 4 5 6 7 8 42 Baccaurea lanceolata 0.06 2.00 0.020 0.349 0.402 0.115 0.867 43 Baccaurea sp. 0.05 1.50 0.010 0.262 0.201 0.085 0.548 44 Blumeodendron elatriospermum 0.03 0.50 0.005 0.087 0.101 0.050 0.238 45 Blumeodendron tokbraii 0.20 1.00 0.010 0.175 0.201 0.362 0.738 46 Bridelia glauca 0.06 0.50 0.005 0.087 0.101 0.116 0.304 47 Cleistanthus bakonensis 0.45 14.00 0.100 2.445 2.012 0.820 5.278 48 Drypetes longifolia 0.46 3.00 0.030 0.524 0.604 0.835 1.962 49 Endospermum diadenum 1.28 16.00 0.125 2.795 2.515 2.312 7.622 50 Macaranga diepenhorstii 1.67 34.00 0.170 5.939 3.421 3.012 12.371 51 Mallotus penangensis 0.61 12.00 0.080 2.096 1.610 1.109 4.815 52 Mallotus sp1. 0.01 0.50 0.005 0.087 0.101 0.019 0.207 53 Sapium baccatum 0.73 19.00 0.135 3.319 2.716 1.327 7.362 54 Triginostemon serratus 0.01 0.50 0.005 0.087 0.101 0.026 0.214 11. Fabaceae: 0.52 11.00 0.100 1.921 2.012 0.940 4.873 55 Parkia timoriana 0.11 1.50 0.015 0.262 0.302 0.207 0.771 56 Pithecellobium cf. bubalinum 0.40 9.00 0.080 1.572 1.610 0.717 3.898 57 Sindora leiocarpa 0.01 0.50 0.005 0.087 0.101 0.016 0.204 12. Fagaceae: 2.82 10.50 0.100 1.834 2.012 5.099 8.945 58 Lithocarpus urceolaris 2.31 3.50 0.035 0.611 0.704 4.166 5.482 59 Lithocarpus wrayii 0.50 6.50 0.060 1.135 1.207 0.901 3.244 60 Quercus argentata 0.02 0.50 0.005 0.087 0.101 0.032 0.220 13. Flacourtiaceae: 1.87 22.50 0.215 3.930 4.326 3.373 11.629 61 Hydnocarpus kunstleri 0.61 2.50 0.025 0.437 0.503 1.106 2.045 62 Osmelia maingayi 0.38 5.00 0.050 0.873 1.006 0.682 2.562 63 Pangium edule 0.71 12.00 0.110 2.096 2.213 1.276 5.586 64 Scolopia macrophylla 0.17 3.00 0.030 0.524 0.604 0.309 1.436 14. Icacinaceae: 0.16 3.00 0.03 0.524 0.604 0.282 1.409 65 Stemonurus secundiflorus 0.16 3.00 0.030 0.524 0.604 0.282 1.409 15. Juglandaceae: 0.14 0.50 0.005 0.087 0.101 0.252 0.440 66 Engelhardtia spicata Blume 0.14 0.50 0.005 0.087 0.101 0.252 0.440 16. Lauraceae: 2.66 48.00 0.390 8.384 7.847 4.799 21.030 67 Alseodaphne cf. elmeri 0.15 2.00 0.020 0.349 0.402 0.268 1.019 68 Alseodaphne crassifolia 0.05 1.50 0.015 0.262 0.302 0.092 0.656 69 Cinnamomum iners 1.25 21.00 0.170 3.668 3.421 2.267 9.355 70 Cinnamomum subterapterum 0.02 0.50 0.005 0.087 0.101 0.037 0.225 71 Cryptocarya crassinervia 0.02 0.50 0.005 0.087 0.101 0.028 0.216 72 Endiandra rubescens 0.08 2.00 0.020 0.349 0.402 0.142 0.894 73 Litsea glutinosa 0.11 3.50 0.020 0.611 0.402 0.200 1.214 74 Litsea noronhae 0.92 15.00 0.115 2.620 2.314 1.653 6.587 75 Litsea sp1. 0.05 1.50 0.015 0.262 0.302 0.093 0.657 76 Litsea sp3. 0.01 0.50 0.005 0.087 0.101 0.019 0.207 17. Lecythidaceae: 0.02 1.00 0.010 0.175 0.201 0.033 0.409 77 Barringtonia macrostachya 0.01 0.50 0.005 0.087 0.101 0.016 0.204 78 Barringtonia scortechinii 0.01 0.50 0.005 0.087 0.101 0.017 0.205 18. Leeaceae: 0.01 0.50 0.005 0.087 0.101 0.017 0.205 79 Leea sp. 0.01 0.50 0.005 0.087 0.101 0.017 0.205 19. Melastomataceae: 0.16 6.00 0.055 1.048 1.107 0.288 2.442 80 Pternandra caerulescens 0.16 6.00 0.055 1.048 1.107 0.288 2.442 20. Meliaceae: 1.47 14.50 0.140 2.533 2.817 2.648 7.998 81 Dysoxylum sp1. 0.85 7.00 0.065 1.223 1.308 1.534 4.065 82 Dysoxylum sp2. 0.19 2.00 0.020 0.349 0.402 0.334 1.086 254 REINWARDTIA [VOL.12 Appendix 1. continued. Family and Species Basal area (m²) Density (trees/ha) Frequency (%) Relative Density (%) Relative Frequency (%) Relative Basal Area (%) Importance Value (%) 1 2 3 4 5 6 7 8 83 Dysoxylum sp3. 0.01 0.50 0.005 0.087 0.101 0.024 0.212 84 Dysoxylum sp4. 0.05 0.50 0.005 0.087 0.101 0.096 0.283 85 Lansium domesticum 0.12 2.50 0.025 0.437 0.503 0.211 1.151 86 Sandoricum koetjape 0.25 2.00 0.020 0.349 0.402 0.449 1.201 21. Moraceae: 1.27 16.50 0.160 2.882 3.219 2.290 8.391 87 Artocarpus elasticus 0.06 0.50 0.005 0.087 0.101 0.110 0.298 88 Artocarpus kemando 0.75 12.00 0.115 2.096 2.314 1.353 5.763 89 Sloetia elongata 0.46 4.00 0.040 0.699 0.805 0.827 2.331 22. Myristicaceae: 1.39 11.50 0.105 2.009 2.113 2.508 6.629 90 Horsfieldia cf. subglobosa 0.28 5.00 0.045 0.873 0.905 0.514 2.293 91 Horsfieldia grandis 0.21 3.00 0.030 0.524 0.604 0.374 1.502 92 Horsfieldia macrocoma 0.54 0.50 0.005 0.087 0.101 0.972 1.160 93 Knema mandaharan 0.35 2.50 0.020 0.437 0.402 0.631 1.470 94 Myristica maxima 0.01 0.50 0.005 0.087 0.101 0.017 0.205 23. Myrsinaceae: 0.59 6.50 0.055 1.135 1.107 1.068 3.310 95 Ardisia fuliginosa 0.01 0.50 0.005 0.087 0.101 0.015 0.203 96 Ardisia lanceolata 0.51 5.50 0.045 0.961 0.905 0.926 2.792 97 Ardisia sp1. 0.07 0.50 0.005 0.087 0.101 0.127 0.315 24. Myrtaceae: 2.28 32.00 0.285 5.590 5.734 4.120 15.444 98 Eugenia acutangulum 1.39 22.50 0.195 3.930 3.924 2.507 10.360 99 Eugenia jamboloides 0.38 4.50 0.040 0.786 0.805 0.691 2.282 100 Eugenia polyantha 0.02 0.50 0.005 0.087 0.101 0.030 0.218 101 Eugenia sp3. 0.04 0.50 0.005 0.087 0.101 0.069 0.257 102 Syzygium laxiflorum 0.07 1.50 0.015 0.262 0.302 0.128 0.691 103 Syzygium racemosum 0.38 2.50 0.025 0.437 0.503 0.694 1.634 25. Olacaceae: 0.95 5.50 0.055 0.961 1.107 1.715 3.783 104 Strombosia javanica 0.95 5.50 0.055 0.961 1.107 1.715 3.783 26. Podocarpaceae: 0.013 0.50 0.005 0.087 0.101 0.023 0.211 105 Podocarpus sp1. 0.013 0.50 0.005 0.087 0.101 0.023 0.211 27. Polygalaceae: 0.29 5.50 0.050 0.961 1.006 0.528 2.495 106 Xanthophyllum affine 0.27 5.00 0.045 0.873 0.905 0.494 2.273 107 Xanthophyllum eurhychum 0.02 0.50 0.005 0.087 0.101 0.034 0.222 28. Proteaceae: 0.04 1.50 0.015 0.262 0.302 0.065 0.629 108 Helicia petiolaris 0.04 1.50 0.015 0.262 0.302 0.065 0.629 29.Rhizophoraceae: 0.03 1.50 0.015 0.262 0.302 0.047 0.611 109 Gynotroches axillaris 0.03 1.50 0.015 0.262 0.302 0.047 0.611 30. Rosaceae: 0.04 0.50 0.005 0.087 0.101 0.067 0.255 110 Parastemon urophyllus 0.04 0.50 0.005 0.087 0.101 0.067 0.255 31. Rubiaceae: 0.25 5.00 0.050 0.873 1.006 0.453 2.333 111 Neonauclea sp. 0.03 1.00 0.010 0.175 0.201 0.051 0.427 112 Plectroniella didyma 0.13 2.00 0.020 0.349 0.402 0.226 0.978 113 Randia macrophylla 0.09 1.50 0.015 0.262 0.302 0.162 0.726 114 Rubiaceae spec1. 0.01 0.50 0.005 0.087 0.101 0.014 0.202 32. Rutaceae: 0.15 3.00 0.030 0.524 0.604 0.273 1.400 115 Clausena engleri 0.08 1.50 0.015 0.262 0.302 0.143 0.707 116 Euodia robusta 0.06 1.00 0.010 0.175 0.201 0.103 0.479 117 Euodia sp1. 0.01 0.50 0.005 0.087 0.101 0.027 0.215 2006] PRIYATNA et.al.: Recovery of lowland Dipterocarp forest 255 Appendix 1. continued. Family and Species Basal area (m²) Density (trees/ha) Frequency (%) Relative Density (%) Relative Frequency (%) Relative Basal Area (%) Importance Value (%) 1 2 3 4 5 6 7 8 33. Sapindaceae: 1.06 13.50 0.145 2.358 2.918 1.922 7.198 118 Elattostachys sp. 0.01 0.50 0.005 0.087 0.101 0.018 0.206 119 Lepisanthes alata 0.16 1.50 0.015 0.262 0.302 0.297 0.861 120 Nephelium lappaceum 0.57 6.00 0.055 1.048 1.107 1.029 3.184 121 Nephelium ramboutanake 0.27 3.50 0.035 0.611 0.704 0.493 1.808 122 Pometia pinnata 0.05 2.00 0.035 0.349 0.704 0.085 1.139 34. Sapotaceae: 1.32 29.50 0.270 5.153 5.433 2.382 12.968 123 Ganua mottleyana 0.68 12.00 0.100 2.096 2.012 1.235 5.343 124 Palaquium dasyphyllum 0.04 1.00 0.010 0.175 0.201 0.070 0.445 125 Palaquium sumatranum 0.47 12.50 0.120 2.183 2.414 0.857 5.455 126 Pouteria malaccensis 0.12 4.00 0.040 0.699 0.805 0.221 1.725 35. Sterculiaceae: 0.12 1.50 0.015 0.262 0.302 0.216 0.780 127 Sterculia cordata 0.03 1.00 0.010 0.175 0.201 0.053 0.429 128 Sterculia oblongata 0.09 0.50 0.005 0.087 0.101 0.163 0.351 36. Symplocaceae: 0.09 3.00 0.030 0.524 0.604 0.160 1.288 129 Symplocos fasciculata 0.09 3.00 0.030 0.524 0.604 0.160 1.288 37. Tiliaceae: 1.15 20.00 0.150 3.493 3.018 2.070 8.581 130 Pentace polyantha 1.15 20.00 0.150 3.493 3.018 2.070 8.581 38. Ulmaceae: 0.09 0.50 0.005 0.087 0.101 0.171 0.359 131 Gironniera subaequalis 0.09 0.50 0.005 0.087 0.101 0.171 0.359 39.Verbenaceae: 0.22 5.00 0.050 0.873 1.006 0.392 2.272 132 Teijsmannoidendron coriaceum 0.21 4.50 0.045 0.786 0.905 0.373 2.064 133 Vitex gamosepala 0.01 0.50 0.005 0.087 0.101 0.020 0.208 TOTAL 55.37 572.50 4.970 100.000 100.000 100.000 300.000 INSTRUCTION TO AUTHORS Taxonomic keys should be prepared using the aligned-couplet type. Manuscripts intended for publication in Reinwardtia should be written either in English, French or German, and represent articles which have not been published in any other journal or proceedings. Each manuscript received will be considered and processed further if it is accompanied by signed statements given independently by two reviewers chosen by the author(s) attesting to its merits as well as its scientific suitability for publication in Reinwardtia. Two printed copies (on A4 paper) of the manuscript of not more than 200 pages should be sent to Editors, together with an electronic copy prepared on Word Processor computer programme using Times New Romance letter type and saved as Rich Text File must be submitted. For the style of presentation authors should follow the latest issue of Reinwardtia very closely. Title of the article should be followed by author's name and mailing address and a one-paragraphed abstract in English (with French or German abstract for papers in French or German) of not more than 250 words. Keywords should be given below each abstract. On a separate paper author(s) should prepare the preferred running title of the article submitted. Strict adherence to the International Code of Botanical Nomenclature is observed, so that taxonomic and nomenclatural novelties should be clearly shown, Latin description for new taxon proposed should be provided, and the herbaria where type specimens are deposited should be indicated. Synonyms should be presented in the long form [name of taxon, author's name, year of publication, abbreviated journal or book title, volume (number): [page]. Maps, line drawing illustrations or photographs preferably should be prepared in landscape presentation to occupy two columns. Illustrations must be submitted as original art accompanying, but separate from, the manuscripts. On electronic copy, the illustrations should be saved in .jpg or .gif format. Legends for illustrations must be submitted separately at the end of the manuscript. Bibliography, list of literature cited or references follow the Harvard System. For each paper published author(s) will receive 25 copies of reprints free of charge. Any additional copies should be ordered in advance and the author(s) will be charged accordingly. ISSN 0034 - 365 X REINWARDTIA Vol. 12. No. 3. 2006 CONTENTS Page BENITO C. TAN, BOON-CHUAN HO, VIRGILIO LINK, EKA A.P. ISKANDAR, IPAH NURHASANAH, LIA DAMAYANTI, SRI MULYATI and IDA HAERIDA. Mosses of Gunung Halimun National Park, West Java, Indonesia ,.... 205 S. DEWI. Stachylidium pallidum Dewi sp. nov. from Java 215 W.J.J.O. DE WILDE, B.E.E. DUYFJES and R.W.J.M. VAN DER HAM. Anangia, a new monotypic genus of Cucurbitaceae from East Mollucas 219 TOPIK HIDAYAT, TOMOHISA YUKAWA and MOTOMIITO. Evolutionary analysis of pollinaria morphology of subtnbe Aeridinae (Orchidaceae) 223 DOLLY PRIATNA, KUSWATA KARTAWINATA and ROCHADI ABDULHADI. Recovery of a lowland dipterocarp forest twenty two years after selective logging at Sekundur, Gunung Leuser National Park, North Sumatra, Indonesia 237 MARTHEN T. LASUT. A new species of Ischaemum from Sulawesi 257 HERBARIUM BOGORIENSE BIDANG BOTANI PUSAT PENELITIAN BIOLOGI - LIPI BOGOR, INDONESIA HALDEPAN 48-100-2-PB RECOVERY OF A LOWLAND DIPTEROCARP FOREST TWENTY TWO YEARS AF Herbarium Bogoriense, Pusat Penelitian Biologi, LIPI, Jl. Ju ABSTRACT ABSTRAK INTRODUCTION STUDY AREA AND METHOD A permanent plot was subjectively established in a selectively logged-over forest in 1982 by Abdulhadi et al. (1987). The plot was two hectares (100 x 200 m) and was divided into In February-March 2000 and August-November 2000 the above tw The density, frequency, and dominance as measured by basal a Composition Gaps Changes in tree density and species richness CONCLUSION ACKNOWLEDGEMENT REFERENCES Palaquium dasyphyllum halbel