PERTAniHfi. Vol.1 No. 2 December Contents Page

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1 PERTAniHfi Vol. No. 2 December 978 Contents Page Preliminary Studies on the Performance of Landrace, Duroc and their Crossbreds in a Tropical Environment K.K. Kuan and T.K. Mak 62 Pupal Distribution of Dacus dorsalis Hendel in Relation to Host Plants and its Pupation Depth Yusof Ibrahim and Rosli Mohamad. 66 Observations on Growth and early Production of some Durian (Durio zibethinus Murr) Clones at Universiti Pertanian Malaysia Orchard Othtnan Yaacob, Mohd Noor Ismail and Awaluddin Hj. Talib. 7 Evaluation of some Long Bean Lines using Early and Late produced Seeds in Two Soil Types T.C. Yap and K.T. Cheok. 82 Tandatanda genetik Orang Asli Semenanjung Malaysia dan Bangsabangsa Bumiputera Sabah, Sarawak dan Brunei. Kumpulankumpulan darah dalam tiga bangsa terbesar di Malaysia dan Singapura. Suatu Penyusunan data S.G. Tan. 86 Effects of Selection for High LitterWeight on Reproductive Performance in Mice Baharin Kassim and R.G. Beilharz. 97 Tax Burden on Rubber, Coconut and Pineapple Smallholders in Johore Mohammed Ariff Hussein. 5 The Floristic Components of the Ground Flora of a Tropical Lowland Rain Forest at Gunung Mulu National Park Sarawak Ruth Kiew. 2 Leg Paralysis Inducement by Risella 7 oil in Lucilia sertcate (Meig) as affected by sex, time, application site, and oil volume G.S. Lim. 2 Progress of Natural Regeneration after Final Felling under the current Silvicultural Practices in Matang Mangrove Reserve P.B.L. Srivastava and Daud Khamis. 26 Mineralogy and Related Properties of Acid Sulphate Soil from Melaka, Peninsular Malaysia Shamsuddin jfusop and Tuan Omar Raja Senik. 36 Short Communication Kesan Nukleotid dan Nukleosid terhadap Aktiviti Fosfodiesterase Mycobacterium smegmatis C.H. Lee. 43 A Scientific Journal published by UNIVERSITI PERTANIAN MALAYSIA KDN 579/78 ISSN 26628

2 f*.*'. :V ARCHIVE COPY (Please Do Not Remove) EDITORIAL BOARD J. J. AUGUSTIN AHMAD MAHDZAN AYOB AZIMI Hj, HAMZAH BADRI MUHAMMAD BAHARIN KASSIM (Chief Editor) (Business Manager) CHIN HOONG FOONG KHAMIS AWANG KWOK CHEE YAN CH'NG GUAN CHOO PERTANIKA is a scientific journal published twice a year by Universiti Pertanian Malaysia and is centred around studies that are related to Agriculture, Forestry, Veterinary Science, Fisheries, Agricultural Engineering, Food Science and Technology, the Basic Sciences, Agricultural Economics, Extension and Continuing Education, and Rural Social Studies. PERTANIKA welcomes original reports, in English or Bahasa Malaysia, of research not previously or simultaneously published in any scientific or technical journal, from the Staff of Universiti Pertanian Malaysia and other local and overseas institutions and organisations. NOTES FOR CONTRIBUTORS appear on the back cover of every issue of the Journal. Contributions are reviewed by a panel of consultants whose names appear in the last issue of each volume. Articles should be submitted to the Chief Editor, PERTANIKA, Universiti Pertanian Malaysia, Serdang, Selangor, Malaysia. Subscription Rates Individuals Institutions Malaysia/Singapore one year M $. M $2. Overseas one year US$5. US$. Overseas subscribers please add M $5. (US $2.) per issue for airmail surcharge. Cheques/Bank drafts should be made payable to UNIVERSITI PERTANIAN MALAYSIA and returned to the Business Manager, PERTANIKA, Universiti Pertanian Malaysia, Serdang, Selangor, Malaysia.

3 PCRTAfllKA CONTENTS OF VOLUME. Number, July 978 Katakata Aluan Professor Tan Sri (Dr) Mohd Rashdan bin Haji Baba Further Studies on the Reestablished Populations Derived from Chinta and the New Sweet Corn Composite T.C. Yap, S.H. Tan and S.T. Tan 3 Use of Cassava and Winged Bean Flour in BreadMaking Ng Eng Gitn and Khor Geok Lin. 7 Relationship of Stalk Strength and Grain Yield in Maize (Zea mays L) Determined by Si Progeny Tests Mohamad Idris Haji Zainal Abidin. 7 TandaTanda Genetik Biokimia dalam tiga bangsa terbesar di Semenanjung Malaysia dan Singapura: Satu Penyusunan Data S.G. Tan. 22 Soil Application of Granular Carbofuran to Control Bean Fly, Ophiomyia phaesoli (Tyron) Mohd Yusof Hussein. 36 Bacterial Fermentation of High Alkaline Waste from Irish Potatoes Mohamed Ismail Abdul Karim. 4 Digestibility of Napier Grass (Pennisetum purpureum) in Grass Carp (Ctenopharyngodon idella) A.T. Law. 5 The Life Cycle of The Fruit Fly, Dacus dorsalis Hendel, on Chilli Fruits Abdul Ghani Ibrahim. 55 Short Communication A Technique for Pollen Studies Lim Eng Siong. 59

4 Number 2, December 978 Page Preliminary Studies on the Performance of Landrace, Duroc and their Crossbreds in a Tropical Environment K.K. Kuan and T.K. Mak 62 Pupal Distribution of Dacus dorsalis Hendel in Relation to Host Plants and its Pupation Depth Yusof Ibrahim and Rosli Mohamad. 66 Observations on Growth and Early Production of some Durian (Durio zibethinus Murr) Clones at Universiti Pcrtanian Malaysia Orchard Othman Yaacob, Mohd Noor Ismail and Awaluddin Hj. Talib. 7 Evaluation of some Long Bean Lines using Early and Late produced Seeds in Two Soil Types T.C. Yap and K.T. Cheok. 82 Tandatanda genttik dalam Orang Asli Semenanjung Malaysia dan Bangsabangsa Bumiputera Sabah, Sarawak dan Brunei. Kumpulan darah dalam tiga bangsa terbesar di Malaysia dan Singapura. Suatu Penyusunan data S.G. Tan. 86 Effects of Selection for High LitterWeight on Reproductive Performance in Mice Baharin Kassim and R.G. Beilharz. 97 Tax Burden on Rubber, Coconut and Pineapple Smallholders in Johore Mohammed Ariff Hussein. 5 The Ploristic Components of the Ground Flora of a Tropical Lowland Rain Forest at Gunung Mulu National Park Sarawak Ruth Kiew. 2 Leg Paralysis Inducement by Risella 7 oil in Lucilia sericate (Meig) as affected by sex, time, application site, and oil volume G.S. Lim. 2 Progress of Natural Regeneration after Final Felling under the current Silvicultural Practices in Matang Mangrove Reserve P.B.L. Srivastava and Baud Khamis. 26 Mineralogy and Related Properties of Acid Sulphate Soil from Melaka, Peninsular Malaysia Shamsuddin Jusop and Tuan Omar Raja Senik. 36 Short Communication Kesan Nukleotid dan Nukleosid terhadap Aktiviti Fosfodiesterase Mycobacterium smegmatis C.H. Lee. 43

5 Rewawh Mamgemtnt Ctn*» (RMC) ntroorioeahii Pertanika (2), 6265 (978) UniveraWPutiiMtftyiia ^4S«fdanQ t S«Uiogof,M«l«yiia Tel: , W4769, Preliminary Studies on the Performance of Landrace, Duroc and their Crossbreds in a Tropical Environment K. K. KUAN and T. K. MAK Faculty of Veterinary Medicine and Animal Science, Universiti Pertanian Malaysia Key words: Pigs, Performance, Repeatability RINGKASAN Prestasi bagi babi di Unit Babi, Universiti Pertanian Malaysia telah dikaji. Berat waktu lahir setiap ekor anak babi bagi kacukan LandraceDuroc lebih tinggi (P<.) daripada berat Landrace dan Duroc di dalam kelahiran pertama dan kedua. Kacukan LandraceDuroc telah pun menunjukkan kemajuan bagi bilangan anak babi pada kelahiran, bilangan anak babi pada masa berhenti menyusu, berat pada masa berhenti menyusu, umur pada estrus pertama, dan umur sewaktu melahiranak pertama. Berat pada masa berhenti menyusu di dalam kelahiran kedua adalah tidak sebegitu baik, kerana prestasi bagi kacukan LandraceDuroc adalah kurang daripada Landrace dan Duroc. Kekurangan ini mungkin disebabkan oleh pertukaran makanan babi. Anggaran perulangan bagi bilangan babi pada kelahiran untuk Landrace dan Duroc adalah.46 dan.246. SUMMARY A study on the performance of pigs at the Pig Unit, Universiti Pertanian Malaysia, was made. The birth weight per pigling of LandraceDuroc crossbreds was significantly higher (P<.) than the mean of the parental breeds in the first and second farrowing. The crossbreds also showed general improvement in litter size at birth, litter size at weaning, weaning weight, age at first estrus and age at first farrowing. The only exception was the weaning weight at second farrowing in the crossbreds where their performance was lower than the purebreds. This could be attributed to a change of feed. Repeatability estimates of.46 and.246 for litter size at birth in Landrace and Duroc were obtained. INTRODUCTION In recent years the trend of swine production in Malaysia has been towards the use of exotic breeds and their crosses. Mahendranathan (97) has estimated that the Local Chinese Pig accounts for less than.5% of the local pig population. In the same paper the reasons for the decline in the popularity of the Local Chinese pig were enumerated. Currently, the popular exotic breeds of pigs in the country are Landrace, Duroc, Yorkshire, Hampshire, Chesterwhite and their crosses. Research data on the reproductive and general performance of these breeds are, however, limited. The object of this study was to document some of the reproductive and performance data of two of these popular exotic breeds and their crosses. The observations in this study were made at the Pig Unit at Universiti Pertanian Malaysia in Puchong, about 8 miles from the main Campus. The Pig Unit is located on a hill slope at the 62 fringe of a jungle reserve with a good running stream. Built in 975, it initially comprised a breeder barn, a farrowing house and nursery house. In 977, a growingfinishing house was added to the Unit. The farm today has a capacity for 4 sows and 25 growingfinishing pigs. MATERIALS AND METHODS The farm came into operation in early 976. Initially, 3 Duroc and Landrace gilts together with two boars of each breed were purchased from the Government Multiplication Station at Serdang. The animals were acquired when they were between three to five months of age. The crossbreds are the progenies of the original stocks. The current practice at the farm is to keep gilts in pens with four animals to a pen and the boars in individual pens next to the gilts. The object we of this arrangement is to stimulate the gilts to come to early maturity. Feeding is on an ad lib. basis with a proprietory grower's ration of 6% crude protein for both the gilts and boars.

6 When the animals reach a liveweight of about 9 kg, feed intake is restricted to 2.3 kg a day, twice daily, to avoid excessive fatness. Water is available at all times through automatic water nipples. Careful observations are made as to periods the gilts come into heat and the dates are recorded. Gilts which come into first heat at between five to six and a half months of age, have their mating delayed until they are between seven to eight months of age. It is hoped that the delayed mating will result in a larger litter size as the number of ova shed per estrus increases gradually over the first several estrus cycles (Pond and Maner, 974). Gilts which come into their first estrus after seven months are, however, bred at their first heat since it is not economical to wait longer than this. Matings are by natural service. Gilts of one breed are mated to boars of the other breed i.e. Duroc gilts are mated to Landrace boars and vice versa. Gilts are mated a second time 224 hours after the first mating as this practice is associated with a higher conception rate resulting in one or two extra pigs per litter. Once the gilts have been bred, they are restricted to 2.3 kg of a commercial breeder mash and put under observation to see whether they come into heat again 924 days later. When pregnancy is confirmed the animals are moved into gestation stalls and kept there until about one week before farrowing. The animals are then moved out of the gestation stalls, thoroughly scrubed and moved into the farrowing stalls in the farrowing house. K. K. KUAN AND T. K. MAK A livestockman is present at all farrowings. When the piglings are born their needle teeth are clipped, their ears are notched for identification and they are weighed within six hours of birth. At three days of age an intramuscular injection of mg iron dextran is given into the ham muscles. Male piglings are castrated at two weeks of age. Piglings are weighed at weekly intervals. The daily amount of feed given to the Iactating sow is based on the number of piglings she produces. Each sow receives a maintenance ration of.36 kg plus.45 kg for every pigling she has. For example, if a sow has given birth to eight piglings she receives 4.9 kg of a breeder ration per day. Piglings are introduced to creep feeding at ten days of age. Weaning is at five weeks. All deaths and their causes are recorded. Ten good gilts based on general appearance, growth rate and feed efficiency were selected from this first mating for our own farm use. Their management is similar to that described for the pure breds. Mating of the crossbred gilts is by the crisscross method. Observations in this study were made under the management conditions described above. RESULTS Table shows the mean values of litter size at birth, birth weight per pigling, weaning weight per pigling and number of piglings weaned. The LandraceDuroc crossbreds were found to have bigger litter size compared to the Landrace and Duroc in the first farrowing. The same trend was seen in the second farrowing. However, the differences among the various types of pigs were not significant in both the first and the second farrowing. st Farrowing 2nd Farrowing Type of Pigs Landrace Duroc LandraceDuroc crossbred Landrace Duroc LandraceDuroc crossbred TABLE Mean Values of Litter Size at Birth, Weaning Weight Per Pigling and No. of Sows Litter Size at Birth 8. ± ± ± ± ± ±.9 63 Birth Weight Per Pigling, Litter Size at Weaning Birth Wt. per Pigling (kg).34 ±.6.4 ±.5.47 ±.46.3 ±.7.22 ±.6.53 ±.6 Weaning Wt. per Pigling (kg) 9.3 ± ± ± ± ± ±.4 Litter Size at Weaning 7.22 ± ± ± ± ± ±.9

7 PERFORMANCE OF PIGS IN TROPICAL ENVIRONMENT The birth weight of the crossbreds differed at a highly significant level from that of the Duroc as well as the Landrace and Duroc combined (P<.) in the first farrownig. The amount of heterosis, estimated by comparing the mean of the crossbreds with that of the parents, was 2.5%. In the second farrowing, the birth weight of the crossbreds was also significantly different from that of the Landrace (P<.5) the Duroc, as well as the Landrace and Duroc combined (P<.), while the heterotic advantage was 2.68%. Weaning weight in crossbreds in the first farrowing was higher than that of the Duroc, and the Landrace and Duroc combined, but was lower than that of the Landrace. These differences, however, were not significant. In the second farrowing the weaning weight of the crossbreds was significantly lower than that of the Duroc, and the Landrace and Duroc combined (P<.5). This significant difference may be due to the change of feed given to the Duroc, which caused them to show unusual increase in weaning weight. The crossbreds showed a higher number of piglets weaned when compared to the Landrace and Duroc in the first farrowing. In the second farrowing the number of piglets weaned was slightly higher than the mean of the Landrace and Duroc. In both cases there were no significant differences. As shown in Table 2, the crossbreds exhibited a lower age at first estrus and at first farrowing; the Duroc was intermediate in values in these two characteristics studied. The differences were, however, not significant. The repeatability estimate of litter size as calculated according to the method by Becker (975) is shown in Table 3. Duroc showed a higher repeatability estimate than Landrace. DISCUSSION A litter size at birth of 7.8 and 8. in Landrace in the first and second farrowing respectively Landrace Duroc TABLE 3 Repeatability of litter size at birth No. of sows 7 7 No. of litters Repeatability.46 ± ±.235 was reported by Mahendranathan and Mellish (972). These values are lower than those shown in Table for Landrace. Anuwar (968) reported the litter size for Landrace to be 8.9 in the first farrowing. This is in agreement with the value of 8. obtained by us. However, his observation of 7.85 in the second farrowing is much lower than our value of 9.29 for Landrace (Table ). In the crossbreds (different types), Selvarajah (967) found the litter size to be 8.2 and 9.6 for the first and second farrowing respectively while Mahendranathan and Mellish (972) obtained values of 8.9 and 9.6 respectively on commercial farms. The LandraceDuroc crossbreds in our study had a litter size of 9.5 and 9.5 for the first and second farrowing respectively. The three values obtained for litter size in the second farrowing appear to be in general agreement. Mahendranathan and Mellish (972) reported the litter size for Landrace at weaning to be 6.5 and 6.6 in the first and second farrowing respectively; and for the crossbreds, the litter size at weaning was 8.3 in the first farrowing. In our study, the Landrace figures of 7.22 and 8.43 for litter size at weaning in the first and second farrowing were higher than those reported by them, while our crossbreds had a litter size at weaning of 8.88 (Table ). Age at first estrus and age at first farrowing were and days respectively. These figures were lower than those for the purebreds (Table 2). Though the differences were not statistically significant, the early maturity and early farrowing in crossbred sows could be of advantage to commercial pig farmers. Landrace Duroc LandraceDuroc crossbred TABLE 2 Mean values of age at st estrus and age at st farrowing No. of sows Age at st Estrus (days) 28.3 ± ± db 6. Age at st Farrowing (days) ± ± ± 5.24

8 K. K. KUAN AND T. K. MAK The repeatability estimate of.246 for litter size in Duroc appears to be high compared to the Landrace which had a repeatability estimate of.46 (Table 3). The high standard error of repeatability was due to the very small sample size in both cases. Repeatability estimates of litter size in sows vary from.2 to.8 (Pirchner, 969). Turner and Young (969) indicated that repeatability estimates can be used for predicting the increase in lifetime production which can be achieved through early selection. On the basis of the results of this limited study it would appear that crossbred sows generally perform better than purebred ones. ACKNOWLEDGEMENTS We wish to thank the Farm Director Universiti Pertanian Malaysia, for the use of the facilities of the Pig Farm. We also wish to express our appreciation to the former Pig Manager, Mr. Lee Chin Nyean and the Livestockman, Mr. Toh Yan Wing, for their help in the collection of data. REFERENCES ANUWAR BIN MAHMUD (968): Phenotypic and genetic parameters of pigling traits from birth to weaning. Malays. Vet. J. 4: 226. BECKER, W.A. (975): Manual of Quantitative Genetics. 3rd Washington State University Press, Washington. MAHENDRANATHAN, T. (97): The future of the Local Chinese Pigs West Malaysia Industry. pp. Ministry of Agriculture and Fisheries, Kuala Lumpur, Malaysia. MAHENDRANATHAN, T. and MELLISH, K. (972): Some observation on the reproductive performance of Pigs at the Livestock Station, Serdang. Paper presented at the Seminar On Animal Reproduction and Artificial Insemination, Kuala Lumpur. 7th. April, 972. PIRCHNER, FRANZ. (969): Population Genetics in Animal Breeding. San Francisco. W.H. Freeman and Co. POND, W.G. and MANER, J.H. (974): Swine Production in Temperate and Tropical Environments. San Francisco. W.H. Freeman and Co. SELVARAJAH, T. (967): Some gsnetic studies on the litter size of crossbred pigs. Kajian Vet., : TURNER, H.N. and YOUNG, S.S.Y. (969). Quantitative Genetics in Sheep Breeding. Ithaca. Cornell University Prses. (Receivd 2 May 278) 65

9 Pertanika (2), 6669 (978) Pupal Distribution of Dacus dorsalis Hendel in Relation to Host Plants and its Pupation Depth YUSOF IBRAHIM and ROSLI MOHAMAD Department of Plant Protection, Faculty of Agriculture, Universiti Pertanian Malaysia Key words: Fruit fly Dacus dorsalis Hendel; Pupal Distribution; Pupation depth. RINGKASAN Penyelidikan mengenai pertaburan kepompong Dacus dorsalis Hendel berkaitan dengan pokokpokok perumah dan ujian paras dalam kepompongan ditanah telah dijalankan. Sampelsampel daripada pokok betik menunjukkan kelebihan bilangan kepompong (P<..5) berbanding dengan bilangan yang didapati dari pokokpokok jambu, belimbing segi, dan nangka. Tidak terdapat perbezaan diantara ketigatiga pokok perumah yang kemudiannya. Kepompong juga telah didapati bertaburan lebih kilrang samarata dibawah pokokpokok perumah. Keputusan yang didapati juga menunjukkan serangga ini lebih gemar berkepompong pada paras 2 cm dan 3 cm dalam tanah. SUMMARY Studies on pupal distribution of Dacus dorsalis Hendel in relation to host plants and its depth of pupation were conducted. Samples from papaya trees gave significantly (P<.5) higher pupal count compared to those from guava, starfruit, and jackfruit. No significant differences were obtained among samples from the latter three host plants. The pupae were also found to be almost evenly distributed under the trees. The results obtained also showed that this insect prefers to pupate at 2 cm and 3 cm soil depth. INTRODUCTION The oriental fruit fly, Dacus dorsalis Hendel, is an important pest of fruit trees in Malaysai. Starfruit (Averrhoa carambola) is one fo the important hosts of this pest. Other fruits include papaya ( Carica papaya), guava (Psidium guajava) and the jackfruit (Artocarpus heterophyllus). Work on this pest had been reported as early as 93 in the Philippines (Jones), 94 in Indonesia (Dammerman), and 94 in Malaysia (Miller). The females oviposit the eggs immediately beneath the skin of the fruit. The fruit soon becomes damaged as the newly hatched larvae burrow and consume the succulent flesh of the fruit. The affected fruits drop prematurely resulting in poor yield. The lifecycle is completed in the soil where the larva pupates and emerges in ca. days later. Current control practices involve bagging and/or insecticide spraying of the fruits. These procedures, however, are laborious and time consuming. Furthermore, the success of the control depends on the precise timing of the spraying operation* There is also the danger of insecticidal residues being consumed. 66 In view of the above problems a study was conducted to determine the distribution of the insect in the soil, and its behavioural activity with regard to pupation depth. MATERIALS AND METHODS Two related studies were undertaken: a field study on the pupal distribution in relation to host plants; and a laboratory observation on the pupation depth of the pest. Pupal Distribution: Four fruit areas, papaya, starfruit, guava, and jackfruit in the University Farm were selected. Five fruiting trees were selected randomly from each area. Four soil samples per tree spaced 3.5 cm, 6 cm., 9.5 cm., and 22 cm., away from the stem were randomly taken under the canopy where heavy fruit sets were located. Each soil sample, 5.25 X 5.5 X cm. deep (6" x 6" x 3"), was placed in a saturated solution of NaCl to separate the pupae and the debris from the soil. This was then passed

10 YUSOF IBRAHIM AND ROSLI MOHAMAD through a 25 mesh sieve (Endecoff Ltd., London) and allowed to dry for about 2 hr. before the actual pupal count was made. Pupation depth: In this study paper containers as in Fig. were used to determine the pupation depth of D. dorsalis. Petri dishes of 95 cm. diameter were used as the bases. Six layers of Manila cards, each cm. taller th?.n the successive inner layer, were wrapped around each petri dish and secured with cellophane tape. The containers were then filled to the brim with loose, friable, moist soil (Serdang Series: Sandy loam top soil). TABLE Pupal count of Dacus dorsalis Hendel to random samples of fruit trees. No. of Observations Total Mean Papaya A No. of pupae recovered Guava B Starfruit B Jackfruit B Means followed by the same letter are significantly different (P<.5) as determined by DMRT. Figure : Manila card container used in the pupation depth study. Thirty 3rd. instar fruit fly larvae were released into the container which was then immediately covered with another petri dish to prevent larvae from escaping. The cover was removed after all the larvae had successfully burrowed into the soil. Pupation depth, as determined by pupal count made five days later, was recorded by successive removal of the outer wall of the container. The test was replicated eight times at room temperature of ca. 28 C. RESULTS AND DISCUSSIONS Pupal Distribution : Pupae recovered from samples in papaya areas were greater in number (P<.5) than those from guava, starfruit, or jackfruit (Table ). This is perhaps due to the heavy fruiting nature of the papaya trees compared to the other trees sampled. The result also showed no significant 67 difference in pupal number recovered among guava, starfruit and jackfruit. There were no significant differences among all the treatments (Table 2). This indicates that the pupae were randomly distributed under the canopy of all the fruit trees sampled. Pupation Depth: The 2cm and 3cm depth of soil 9ecm to be the preferred regions of pupation. As shown in Table 3, highly significant (P<.) readings were recorded at these depths in comparison to those at the other levels. AliNiazee (974), however, reported from a field study that cherry fruit fly pupae preferred to diapause within the top 4 in (approx. cm) of soil depth. He attributed this preference to the physical and environmental factors of the soil. Shah et al. (948) found that there was variation in the depth of pupation in ploughed and unploughed soils. Cavalloro and Delria (975) believed that chemical factors did not have any influence on the depth of pupation. Their study revealed that the fruit fly Ceratitis capitata pupated deeper in cracked and dry soil than they did in wet soil. In this investigation it would appear that at the regions of 2cm and 3cm soil depths, the soil environment would be most favourable in that it would not affect pupal development and eventual adult emergence. CONCLUSION Significantly more pupae of Dacus dorsalis Hendel were recovered from the papaya trees than from the other fruit trees sampled. The results also indicate no significant differences

11 Sampling dist. (cm) fr. stem Papaya PUPAL DISTRIBUTION OF DACUS DORSAUS TABLE 2 HENDEL Number of pupae recovered at various sampling distances from the host plants. No of pupae recovered per observation Total Mean Gitava Starfruit Jackfruit No significant difference (P>.5) among all treatment means as determined by DMRT Sampling depth (cm) TABLE 3 Pupation depth of Dacus dorsalis HendeL No. of pupae recovered in each replicate Means followed by the same letter are highly significantly different (P<.) as determined by DMRT. among samples from guava, starfruit, and jackfruit trees. The distribution of the pupae under these trees was also about even. Laboratory tests on pupation depths showed that in loose, friable, and moist soil the larvae of the insect preferred to pupate at 2 cm and 3 cm depth of soil ACKNOWLEDGEMENT Mean.88b.3a 8.25a 4.3b.63b.3b We wish to express our sincere thanks to En. Abd. Ghani Ibrahim, Plant Protection Department, Universiti Pertanian Malaysia, for reviewing this manuscript and to all those who helped in this study.

12 YUSOF IBRAHIM AND ROSLI MOHAMAD REFERENCES ALINIAZEE, M.T. (974): The Western cherry fruit fly, Rhagoletis indifferens (Diptera: Tephritidae). Distribution of the diapausing pupae in the soil. Can. Ent. 6:9992. CAVALLORO, R. and DELRIA, G. (975): Soil factors influencing the pupation of Ceratitis capitata Wiedmann. Bull Lab. Ent. agra. 'Filippo Silvestri* Portici. 32: 995. (Abstract) DAMMERMAN, K.W. (94): The fruit fly question in Java Meded. v.d. Afdeel. v. Plantenziekten, Dept. van Landboun, Nijverheid en Buttenzorg, No. 8, pp. 2 (Abstract). Handel, JONES, CR. (93): The Mango fruit fly and other pests in the Philippines. Philipp. agric Rev. Part, 3: 442 (Abstract). MILLER, N.C.E. (94): 28:22. Fruit flies. Malay. Agric. J. SHAH, M.L, BATRA, H.N., and RENJHEN, P.C. (948): Notes on the biology of Dacus (Strttmeta) ferrungineus Fabr. and other fruit flies in Northeast Frontier Province. ind.j. Ent. : (Received 7 September 978) 69

13 Pertanika (2), 78 (978) Observations on Growth and early Production of some Durian (Durio zibethinus Murr) Clones at Universiti Pertanian Malaysia Orchard OTHMAN YAACOB, MOHD. NOOR ISMAIL and AWALUDDIN HAJI TALIB 2 Department of Soil Science, Faculty of Agriculture, Universiti Pertanian Malaysia Key words: Durian growth, Early yield, Research needs. RINGKASAN Penyata mengenai tanaman durian diatas kawasan tanah bekas getah tua seluas 22 ekar adalah dibentangkan bersama dengan rekodrekod sukatan hujan, kadar membaja dan Iainlain kawalan antali yang dibuat dalam kawasan itu semenjak 969. Kadar tumbohsaran semua jenis klon durian yang ditanam, keadaan membunga dan membuah serta dengan sebabsebab kekurangan buahbuah bagi klon D2 dan D96 juga di bincangkan. Dengan adanya keadaan saperti itu, penyelidikan untuk mengatasi setengah daripada masalah masalah tertentu telah pun dicadangkan untuk masa hadapan. SUMMARY The establishment of a 22 acre durian orchard on an oxisol soil formerly under old rubber is recorded outlining the nature of the environment and the cultural practices adopted since 969. The growth pattern of all clones planted, early flowering and fruiting patterns are also discussed together with possible causes of poor fruiting habits of D2 and D96. Some possible areas for research are suggested for future work based on these observations. INTRODUCTION The object of this paper is to provide background information which may be required for any future detailed study on durian at Serdang. The 22acre site under study was at one time a part of the Air Hitam Forest Reserve. It was cleared for planting with rubber probably in the 94's by Serdang Estate which at that time owned most of the land on which the University Farm is now situated. The area was purchased with other land by the University in 966; and in late 968 it was mechanically cleared. The trees were felled and the trunks were sawn up for wood and removed from the site. The remaining roots and stumps were collected into large heaps and burnt after which the area was ploughed and rotovated three times. Remaining roots were hand removed and burnt; and after a short period of dry weather, planting holes measuring 2' X 2' X 2' and spaced 4' x 4' apart were prepared. This spacing allowed for 27 trees to be planted per acre. In each planting hole about 2 pounds of rotted cattle and poultry manure were placed three weeks prior to planting. In July 969, during the wet season, the recommended durian clones were planted. During the first four years, watering was carried out during dry periods, or as and when necessary. The general maintenance consisted of regular weeding and spraying against pests and diseases, carried out by the Farm Division and by the Plant Protection Department respectively. DURIAN CLONES PLANTED AND FERTILIZER PROGRAMME USED Durian clones The known clones planted were D2, D7, D8, D, D24, D66, D84, D88, D96, and an unidentified clone designated as DR. The num Farm Director, Universiti Pertanian Malaysia. 2 Senior Field Assistant, Universiti Pertanian Malaysia. 7

14 OTHMAN YAACOB, MOHD. NOOR ISMAIL AND AWALUDDN HAJI TALIB ber of trees planted for each clone and the percentage composition of each clone in the area is given in Table. The actual distribution on the ground is shown in Fig.. From the time planting began in 969 until 973, virtually all clones were replaced at various growth stages, mainly because of disease. Clone D2 D7 D8 D D24 D66 D84 D88 D96 DR TABLE Durian clones planted at the Universiti Pertanian Malaysia Orchard, Serdang. Number of trees planted Percentage of total stand Total 578. Fertiliser programme The fertiliser programme followed the 'standard' practice recommended by Whitehead (959). The rates of application over the years are shown in Table 2. nutrient content decreasing with depth. At the intermediate site, however, the nutrient content is high in the upper layer 6 cm deep, where most of the feeding roots are located. In the valley site the soil is relatively lighter in texture with nutrient contents comparable to the hill site (Table 3). This is somewhat unexpected as soils on the lower sites normally have high nutrient contents. It is probable that during the eight year period, (969978), the nutrients applied may have leached down the slope as suggested by the high nutrient contents of soil at the intermediate site. Details of this possibility occurring are being studied as part of the current research on durian. Rainfall The monthly and annual rainfall at Serdang from 968 to 977 are given in Table 4. The average rainfall for the year period (969977) was 23 mm although the annual rainfall for 974 and 975 was below average. The two dry years adversely affected the general yield of durian throughout the country (Anon 976). * AGRONOMIC OBSERVATIONS Growth pattern There are several parameters by which growth can be measured. In this study, growth was measured by taking the height of the tree from the base of the trunk at ground level to the top of the canopy, and the total spread of the canopy from left to right. A specially constructed light alluminium frame was used for this purpose (Plate ). Topography and soils NATURE OF SOILS AND THE ENVIRONMENT The orchard can be divided into two broad sites based on the general topography of the area and these are conveniently designated as the Valley' and the ( hih\ A rise resulting in a slope of more than is designated the 'hill and a slope below this angle is designated the 'vallev' (Fig. 2). The mechanical and chemical composition of the two soil types found in the designated sites are given in Table 3. A portion of the valley is made up of collovial soil while the hill is almost entirely of Malacca soil with a varying depth of lateritic layers, the 7 The growth shape of durian trees appeared to differ slightly among clones. The average range of height of trees varied from 5.5 for D88 to 7.8 m for the unidentified (DR) clone. The spread of the canopy was narrow in D88, being.8 to.9 m, and wider in D24, from 3. to 3. m. The average height of the remainder of the clones ranged between 5.8 to 7.5 m for height; and 2.3 to 2.4 or 2.9 to 2.8 for canopy spread (Table 5). At Serdang the growth shape characteristics of each clone remain constant (Plates 2a 2e) but these characteristics vary slightly in other parts of the country because of different environmental conditions. However, no information on this variable characteristic is available at the present. Flowering pattern First flowering of certain clones was observed in September 974. Flowering from all trees

15 <X^ o a 7 o R L i E A R C H ( LOT > m > r 7 O DC. " \ \ \ O o D C 2 V n r O m * { i. Jo?c Z:a Co ^5 "?.7.2 : Tractor hath. * ^ ^ [.^, ^d u 38 Od D8 D3 ^88 D84 D66/9 6 D8 Fig. Distribution of durian clones planted on the Universiti Pertanian orchard.

16 OTHMAN YAACOB, MOHD. NOOR ISMAIL AND AWALUDDIN HAJI TALIB TABLE 2 Fertiliser programme and rate (oz/tree) for durian at the Universiti Pertanian Malaysia: Year 969 Season/month of application July September December Fertiliser type*/ grade CCM 2 CCM 2 CCM 2 Amount/rate (oz/tree) January March June July September December Nitro 26 %N CCM 24 CCM 22 Nitro 26%N CCM 22 CCM January March June September December Nitro 26 /o N CCM 44 CCM 22 CQM 22 CCM January June December Nitro 26%N CCM 66 CCM April September CCM 22 CCM April September December CCM 25 CCM 25 CCM April September December CCM 25 CCM 25 CCM September CCM July October Muriate of Potash CIRP February CCM 44 * The range of compound formulations in the ferilisers used is given below: CCM 2 CCM 22 CCM 24 CCM 25 CCM 44 CCM 66 Nitro26 Fertiliser compound no. Christmas Is. (CIRP) Rock Phosphate m Muriate of Potash % N % P2O % K2O % MgO * Total nutrients J 43 42

17 O o H X D m 7 O D n o o Collovium, good drainage, laterite > 5cm Malacca soil laterite >5cm laterite 255cm laterite 25cm Pond/stream D C n O m Building Fig. 2 Soil map of the orchard.

18 OTHMAN Site Mechanical and chemical composition of soils at three sites located on the 22acre orchard at the University Orchard Depth (cm) 'Hill' 'Intermediate* 3 36 'Valley' CS FS Mechanical S analysis C TABLE : ph 4, P Nutrients Available (ppm) K Ca Mg N Total (%) C OM > COB, MO p Z Q o 73 IAIL AND i s> r C D D N HAJI TALIB

19 GROWTH AND EARLY PRODUCTION OF DURIAN CLONES TABLE 4 Year The monthly and annual rainfall: (mm) at Serdang: J F M A M J J (mm) A s O N D Total yr. average Clone TABLE 5 Growth pattern of durian clones at the Universiti Pertanian Malaysia Orchard No. of trees measured Height Range for Canopy Left Right Height Mean Left Canopy * Right D 2 D 8 D 24 D 66 D 88 D 96 DR Other clones were not measured as they formed only a small percentage of the total (Table ) , beginning in February 977 was subsequently recorded and expressed as a percentage of the total number of trees for each clone. Numbers of fruits obtained were also recorded (Table 6). It is clear that under the environmental conditions recorded and the cultural and manual practices adopted at Serdang, virtually all clones derived from budded planting materials flowered five years after planting. Except for D2 and D88, the number of clones flowering later than five years of age was less than fifteen per cent for the first four seasons in 974 and 975 (Table 6). 76 In 975, the year following the drought when the annual rainfall recorded for 974 and 975 was below the ten years average, three flowering seasons were recorded. The first was in February, the second in May and the third in November. Except for D88, all other clones showed a low percentage of trees flowering in the February fruit season. Clone D2, for example, showed 6% of trees flowering as compared to the 23% recorded for D88 in the May season. In the November season all clones except D88 showed a low percentage of trees flowering with no fruits being recorded at all. Premature dropping and

20 OTHMAN YAACOB. MOHD. NOOR ISMAIL AND AWALUDDIN HAJI TALIB Plate. Technique for measuring the height and canopy spread of a mature durian tree. 77

21 GROWTH AND EARLY PRODUCTION OF DURIAN CLONES a Plate^2a2e Clonal characteristics of some durian trees at the Universiti Pertanian Malaysia Orchard. 78

22 OTHMAN YAACOB, MOHD NOOR ISMAIL AND AWALUDDIN HAJl TALTB theft accounted for the poor fruit recordings. In the following dry year (annual rainfall for 975 was 36 mm), all clones snowed a high percentage of trees flowering in February and July of 976 and some fruits were recorded (Table 6). Observations and recordings indicated that the dry years of 974 and 975 stimulated the production of a large number of fruits throughout the country. It has been suggested that durian It is possible that during a peak period, the developing fruits use up a great deal of stored nutrients; and before another large quantity of fruits can be produced, time is needed to replace the stored materials; this results in high yields occurring in alternate years. The preliminary data on the flowering pattern of the clones appear to support this contention, although it is a too early as yet to confirm this theory. Year and season D2 D8 974 Sept 975 Feb) 975 May) 975 Nov) 976 Feb) 976 Jul) 977 Feb) 977 Jul) TABLE 6 Relationship between the percentage flowered and number of fruits harvested o Flowered D24 D66 D88 D96 D2 D No. of fruits D24 D66 D84 D88 D trees bear fruit biannually; the 975/976 seasons were peak years for the crop (Anon., 976). The peak periods may have coincided with the dry weather as was the case for 975/976 seasons. 79 Clones D2 and D96 did not retain fruits at all under Serdang conditions, although they did well during the early years (948968) at the Federal Experimental Station Serdang (Lee and

23 GROWTH AND EARLY PRODUCTION OF DURIAN CLONES Loh, 966). Fruit drop is also common among various clones. Generally, fruit of the size of golf balls dropped prematurely suggesting that some kind of physiological disorder may have been prevalent to cause premature fruit drop. During the two seasons when flowers and fruit yields were recorded, only D88, which bears largesize fruits weighing between kg, indicated good fruit numbers. The other clones did not give high yields (Table 6). The phenomenon of flower and fruit drop at various stages is being investigated in conjunction with very poor fruiting habits of clones D2 and D96. RESEARCH CONSIDERATIONS Our observations of the performance of durian clones in the Serdang environment suggest that it would be worthwhile undertaking similar studies of other fruit trees grown in the University Farm. Such studies should incorporate: Clonal characteristics and crown shape Although in this study the characteristics of all clones were measured and pictorally recorded (Plates 2a 2e), it would be desirable to extend the research to incorporate a study of the genetic sources of the clones and a detailed morphological study of their differences. Furthermore, research should also include a comparative study of the performance of the same clones in different environments. A comparison of crown shape by the techniques outlined earlier in this paper may require further verification and possible modification for wider use. Furthermore, there is a need for characterisation of individual clones giving details on fruit and vegetative description, supported by pictorial illustration. Other descriptions useful for identification would include enzyme essays, etc. Fertiliser requirements The basic fertilizer requirements of durian, originally a jungle tree, need to be determined if optimum results are to be obtained. Such a study should be extended to include the requirements of other fruit trees. Although in the 22 acre orchard a general fertilizer programme exists (Table 2), a portion of the orchard has been fenced off for detailed experimental research (Fig. ). The type and amount of fertilizers exert a strong influence on the production of fruits. Information on fertiliser effect on durians is limited. Future research should focus on soil 8 fertility, fertilizer recommendation, suitable methods of fertilizer placement, time of application, relationship between fertilizer amount and water regimes on yield and more efficient means of fertilizer application. Flower and fruit drop Despite the high percentage recorded of trees having flowered (Table 6), the fruit retaining ability of almost all clones, particularly D2 and D96, is generally poor. A close look at the planting pattern of durian clones (Figure ) may suggest certain lines of investigation. For example, a large tract of D8, clustered in one area may induce selfincompatibly within the clone and a detailed study on the pollen viability is required. Over and above these, the conspicious absence of arboreal life, so characteristic of a 'dusun' at flowering times, may be a limiting factor in the pollination of durian flowers. For example, bats which pollinate the flowers are virtually absent even at midnight. Bees, another group of efficient pollinators, are also absent as there are no alternative hosts during the durian offseasons. In the absence of such natural pollinating agents, it may be desirable to carry out handpollination similar to that done with oil palm using a few selected trees of all clones; but the economic aspects of adopting this practice need further study. Weeding Under the natural 'dusun' ecosystem, weeding is not carried out except during the fruiting season when the base around each tree is cleared to facilitate easy collection of fruits. However, under an orchard system, weeding with implements driven by heavy tractors is carried out regularly, eventually causing heavy compaction of the top 5 cm soil where the most active feeding root system is located (Othman Yaacob et al. y 977). These procedures affect the surface soil as well as the entire microclimate under each tree. Accordingly, a part of this 22 acre was fenced and used as a research plot (Figure ) where basic quantitive recordings were carried out. Some areas of investigation using this fenced area include: (a) reduction of fertilizer application; (b) reduction of weeding. The effects of these practices on the relative humidity and air temperature in weeded and unweeded plots, with and without fertilizer, are being monitored. As the trees of all clones are now fully developed at seven years of age, and their vegetative growth is satisfactory, fertilizers may not be needed in large quantities. Further

24 OTHMAN YAACOB. MOHD. NOOR ISMAIL AND AWALUDDIN HAJI TALIB more, as the amount required for fruit production is relatively small (Ng and Thamboo, 967) and pruning is minimal, any fertilizer applied would only be for 'fruiting' and not for 'growth'. The effect of fertilizer on young durian trees with and without weeding to approximate 'natural' conditions commonly found in the 'dusun' would be a part of this programme. The effects of these combinations of treatment on the flowering percentage and fruit set are being studied as part of the basic research on durian at the University Farm. ACKNOWLEDGEMENTS We would like to thank all field and laboratory personnel in the Soil Science Department for their technical assistance during the course of this study and Prof. I.C. Enoch for his valuable comments in the preparation of this paper. REFERENCES ANON., (976): "A Feast of Durian Thanks to the Weather/' Malay Mail, 3th July, 976. LEE, C.S. and LOH, CL. (966): Durian varietal trial. Information paper No. 52 Dept. Conf. Min. Agric. Coop. Kuala Lumpur, (cyclostyled memograph). NG, S.K. and THAMBOO, S. (967): Nutrient removal studies on Malaysian fruits: Durian and rambutan. Malay. Agric. J. 46: OTHMAN YAACOB, KHANIF YUSOF and MOKHTARUDDIN MANAN (977): Rainfall pattern and food crop production on oldrubber soils at Serdang. In "Proceedings on FoodAgriculture Malaysia 2". (Ed) H.F. Chin, I.C Enoch and Wan "Mohamad Othman pp.. Faculty of Agriculture, Universiti Pertanian Malaysia. WHITEHEAD, C. (958): The establishment and maintenance of fruit trees. Malay. Agric. J. 42: (Received 9 July 978) 8

25 Pertanika, (2), 8285 (978) Evaluation of Some Long Bean Lines Using Early and Late Produced Seeds in Two Soil Types T. C. YAP and K. T. CHEOK Department of Agronomy and Horticulture, Faculty of Agriculture, Universiti Pertanian Malaysia Key words: Long bean lines, Soil types, Early and Late produced seeds. RINGKASAN Bijibenih kacang panjang yang dihasilkan awal dan lewat dari beberapa keturunan F 9 dari jenis asing telah dinilai di tanah gembur Hat dan tanah lanar di ladang Universiti Pertanian, Serdang. Keputusan percubaan ini menunjukkan hasil kacang dan panjang kacangnya tidak berbeza antara bijibenih dari hasil awal atau lewat. Di antara genotaipgenotaip yang dikaji, hasil kacang tidak berapa tetap jika dibandingkan dengan sifat panjang kacangnya. Setengah keturunan dan jenis Taiwan Stripe Seed memberi penghasilan yang lebih besar berbanding dengan jenis tempatan, iaitu Local Long pada keduadua jenis tanah. SUMMARY Early and late produced seeds of some F 9 lines and exotic cultivars of long beans were evaluated on clayey loam and alluvial soil in the University Farm at Serdang. The results showed that pod yield and pod length were not significantly different for seeds produced early or late. Among the genotypes studied, pod yield was more unstable than pod length. However, some lines and Taiwan Stripe Seed performed better than the local commercial variety, Local Long in both soil types. INTRODUCTION Long bean (Vigna sesquipedalis Fruw.) is also known as asparagus bean or yardlong bean. It is a climbing plant and widely grown in Malaysia for its tender pods. In the past, most long bean varieties grown locally were land strains and were very variable. A breeding programme using different selection procedures was carried out in a 7 X 7 diallel cross population in 973 with the object of developing high yielding varieties. Some of the results for this breeding programme have been reported earlier (Mak, 973, Poh, 976 and Yap et. al, 976). In the present studies some promising F 9 lines of the crop were evaluated under clayey loam and alluvial soil types at the University Farm in Serdang. The present studies also undertook to verify the belief that pods for seeds saved at the early stage of the fruiting period give a better yield than those saved at the later stage. Hence, in addition to the normal genotype evaluation, seeds harvested at the early and late stages of the fruiting period were included. MATERIALS AND METHODS Twelve F9 lines together with three varieties of long bean were used in these studies. Seeds harvested during the early and late stages of the fruiting period were used. The early seeds were pods saved at the beginning of the fruiting period and the late seeds were pods saved 3 days later. Two trials were carried out in these studies. The first consisted of 2 F 9 lines and three varieties were grown on clayey loam in August, 977, while the second was conducted on alluvial soil in September, 977. These genotypes together with the two types of seeds were arranged in a randomized complete block design with three replications. However, due to insufficient seeds, line /5758 was not included in the second trial. The details for the development of the breeding lines have been reported elsewhere (Poh, 976, Yap, etal, 976, 977). The three varieties, Local Long, Taiwan Stripe Seed and Sabah are commercial varieties grown in Peninsular Malaysia, Taiwan and Sabah, respectively. Each genotype was grown on a bed 2 ft (6 cm) long 3.5 ft (7 cm) wide and.5 ft ITemiang Renchong Estate, P.O. Box 3, Panchor, Muar, Johore. 82

26 (5.5 cm) high in two rows spaced 2.5 ft (76 cm) between rows and.5 ft (46 cm) between plants. Two seeds were sown per planting point and later thinned down to one plant per point three weeks after sowing. As long bean is a vine crop, sticks about 8 ft (244 cm) long were set adjacent to each seedling to provide support for the climbing vine. NPK 5:5:5 fertilizer at the rate of 5 kg/ha was applied at the first week and sixth week after sowing. Insecticides were sprayed at weekly intervals to control aphids, bean miners and pod borers. Fresh pods were harvested when they reached marketable stage. Harvesting was carried out on alternate days until senescence of the plants. The two characters reported here were fresh pod yield per plant and pod length. Pod yield per plant was the accumulative yield of fresh pods yield harvested throughout the fruiting period divided by the total number of plants in the bed. Pod length was the mean of pods measured at the third week of harvest. RESULTS Analyses of variance for pod yield and pod length for the two trials are given in Table. Significant differences were found for pod length in the trials but in respect to pod yield only the second trial gave significant differences. Further partitioning of the sum of squares, however, showed that the yield differences among lines in the second trial contributed to the significant difference. In these studies, both trials showed that there was no significant difference in pod yield and pod length in the two types of seeds used for the trials. T. C. YAP AND K. T. CHEOK TABLE There was also no significant interaction effect between genotypes and different types of seeds. Mean values of pod yield and pod length for the two types of the seeds were calculated and compared with Local Long, the commercial variety grown locally. The results showed that Taiwan Stripe Seed and many selected lines outyielded Local Long slightly in both trials. Pod length, however, for most cases was shorter than Local Long (Table 2). DISCUSSION From these studies, it is clear that seed yeild saved at the early and late stages of the fruiting period do not effect any difference in yield and pod length. This finding reveals that farmers could save pods for the next generation at the early or late stage of a fruiting period. It should be noted, however, that pods for seeds should be welldeveloped as is shown by the welldeveloped seeds from good pods used in the present trials. It is generally believed that flowering may be delayed and senescence hastened if mature pods are left unplucked during the fruiting period. If this were the case then it would be logical to keep good pods for seeds at the later stage so that more pods would be harvested for sale. For the two characters studied, both trials showed that pod yield was more unstable than pod length. The correlations between the first and second trials for pod yield and pod length were.3 and.93 respectively, suggesting that pod yield was more susceptible to environmental changes than pod length. These results support the early findings that the heritability for pod yield was Mean squares for characters measured in long bean lines and varieties using early and late produced seeds in two trials Source of variation Degree I of freedom II I Pod yield II I Pod length II Replications ** ** Treatments (29) (27) * 69.82** 74.9** Genotypes (G) * 34.88** 5.44** Stages of seeds (S) G x S Error * P <:.5 ** P <. I : Trial I (clayey loam) II : Trial II (alluvial soil) 83

27 TABLE 2 Mean values of characters measured in long bean lines and varieties using early and late produced seeds in two trials Pod yield/plant (g) Pod length (cm) Variety or line I II I II E L Mean E L Mean E L Mean E L Mean Local Long Taiwan Stripe Seed Sabah r O *»i O o O V435 V6 9 V B H Q 2 r H m on ,4 LSD LSD I : Trial I II : Trial II E : Early L : Late

28 very low compared to that of pod length. Although pod yield of the lines was very unstable under the two soil types, judging by the results of two trials, it was found that some lines performed better than the check variety, Local Long. In these studies, the exotic variety, Taiwan Stripe Seed, also performed quite well under local conditions. The main aim for this breeding programme was to develop high yielding lines; it was found that the yielding ability varied in the two trials. It is necessary for further evaluation trials covering a range of environmental conditions to be carried out to sort out the yielding ability and adaptability of these promising lines. Several experiments have been conducted recently in various regions to evaluate these lines more rigorously. It is hoped that as a result of these experiments some relatively more stable and high yielding lines will emerge and be released for commercial production. ACKNOWLEDGEMENTS This research was supported, in part, by the International Foundation for Science (IFS) in Sweden through the Grant Agreement No. G 2. T. C. YAP AND K. T. CHEOK The authors also wish to express their thanks to Professor Mohd. Zain Karim, Dean of the Faculty of Agriculture, and Dr. Abdul Halim Hassan, Head of the Department of Agronomy and Horticulture, UPM, for their support and guidance in the study. REFERENCES MAK, C. (973): Genetic studies of crude protein content and some agronomic traits in long bean, Vigna sesquipedalis Fruw. M. Agric Sci. thesis. Faculty of Agriculture, University of Malaya, Kuala Lumpur. POH, C.T. (976): Selection studies in early generations of long bean (Vigna sesquipedaiis Fruw.) M. Agric. Sci. thesis. Faculty of Agriculture, University of Malaya, Kuala Lumpur. YAP, T.C., MAK, C. and POH, C.T. (976): Genetic studies and breeding of long beans in Malaysia. Paper presented at National Symposium (SABRAO Malaysia). August 28 29, 976. YAP, T.C., POH, C.T. and MAK, C. (977): Comparative studies on selection methods in long beans. Paper presented in the 3rd International Congress of SABRAO, Canberra, Australia. Feb. 8, 977. (Received 9 September 978) 85

29 Pertanika (2), 8696 (978) Tandatanda Genetik Orang Asli dan BangsaBangsa Bumiputera Sabah, Sarawak dan Brunei. Kumpulankumpulan darah dalam tiga bangsa terbesar di Malaysia dan Singapura: Suatu penyusunan data (Genetic markers in the Aborigines of Peninsular Malaysia and the Indigenous Races of Sabah, Sarawak and Brunei. Blood groups in the three major races of Malaysia and Singapore: A compilation of data). S. G. TAN Jabatan Biologi, Fakulti Sains dan Pengajian Alam Sekitar, Universiti Pertanian Malaysia Key words: Blood groups, Biochemical Genetic Markers, Main Races, Aborigines, Indigenous Races, Malaysia, Singapore, Brunei. RINGKASAN Suatu senarai data berkenaan dengan kumpulankumpulan darah untuk orangorang Melayu, China dan India (tiga bangsa terbesar di Malaysia dan Singapura) dan juga untuk Orang Asli dari Semenanjung Malaysia dan BangsaBangsa Bumiputera dari Sabah, Sarawak dan Brunei, telah disusun dari sastera sains. Data untuk tandatanda genetik biokimia dalam Orang Asli dari Semenanjung Malaysia dan BangsaBangsa Bumiputera dari Sabah, Sarawak dan Brunei pun telah disusun. Taraf adanya data genetik untuk bangsabangsa tersebut dan prospek untuk pengkajian pada tnasa hadapan telah dibincangkan. Kertas ini menyempurnakan suatu percubaan (dalam dua bahagian) untuk menyusun semua data yang ada pada masa sekarang berkenaan dengan tandatanda genetik dalam pendudukpenduduk Malaysia dan Singapura, SUMMARY A list of the data available from the scientific literature on blood groups was compiled for Malays, Chinese and Indians (the three major races of Malaysia and Singapore) as well as for Aborigines from Peninsular Malaysia and the Indigenous Races from Sabah, Sarawak and Brunei. The data for biochemical genetic markers in Aborigines from Peninsular Malaysia and the Indigenous Races from Sabah, Sarawak and Brunei were also compiled. The availability of genetic data for these races and the prospects for further studies are discussed. This paper completes an attempt (in two parts) to compile all the genetic marker data available at present for Malaysians and Singaporeans. Malaysia ialah negara khas sebab populasinya mengandungi orangorang yang berasal dari berbagaibagai bangsa. Sembilan puloh peratus pendudukpenduduk Malaysia berasal dari tiga bangsa terbesar iaitu Melayu, China dan India, dan sepuluh peratus lagi berasal dari kaumkaum Bumiputera lain, terutama sekali kaumkaum Bumiputera dari Sabah dan Sarawak (Rao dll., 977). Di Semenanjung Malaysia, kaum Orang Asli boleh dibahagikan kepada tiga bangsa iaitu Negrito (atau Semang), Senoi dan Melayu Asli. Tiaptiap bangsa itu boleh dibahagikan lagi kepada beberapa suku seperti Semai, Temiar dan Jahut untuk bangsa Senoi dan Jakun, Semelai dan Temuan untuk bangsa Melayu Asli (William Hunt, 952; Carey, 976). Ada kirakira,82 orang Negrito yang tinggal dikawasankawasan 86 terpencil di utara Semenanjung Malaysia, 3,37 Senoi di kawasankawasan pergunungan di pertengahan Semenanjung dan 2,83 Melayu Asli di selatan Semenanjung (Carey, 976). Di Sabah dan Sarawak (dan juga Brunei) selain dari orangorang Melayu dan China, boleh juga dijumpai ramai bangsa Bumiputera yang tidak terdapat di Semenanjung Malaysia. Di Sabah, bangsabangsa Bumiputera terbesar ialah Kadazan (atau Dusun), Murut dan Bajau. Dari jumlah penduduk Sabah (iaitu 84,47), 2,25 ialah orang Kadazan, 35,96 Murut, 9,27 Bajau dan 5,523 orang adalah berasal dari beberapa kumpulan Bumiputera yang lain. Di Sarawak pula, kumpulankumpulan Bumiputera yang terbesar ialah Iban atau Dayak Laut (332,488),

30 S. G. TAN Bidayuh atau Dayak Darat (96,27) dan Melanau (58,734); 56,223 orang berasal dari kaumkaum Bumiputera yang lain. Jumlah penduduk di negeri Sarawak ialah,9,65 (Malaysia 975, Buku Resmi Tahunan). Di Brunei juga, selain dari orangorang Melayu dan China, boleh dijumpai beberapa bangsa Bumiputera seperti orangorang Dayak dan Kedayan. Kertas ini menyempurnakan suatu pcrcubaan untuk mengumpulkan semua data yang ada pada masa sekarang berkenaan dengan tandatanda genetik untuk pendudukpenduduk Malaysia terutama sekali, tetapi termasuk juga data untuk pendudukpenduduk Singapura dan Brunei. Data yang ada untuk tandatanda genetik biokimia bagi tiga bangsa terbesar di Semenanjung Malaysia dan Singapura, iaitu orangorang Melayu, China dan India, sudah pun diterbitkan (Tan, 978). Kertas ini mempersembahkan penyusunan data untuk kumpulankumpulan darah dalam orangorang Melayu, China dan India dari Semenanjung Malaysia dan Singapura dan juga penyusunan data genetik yang ada untuk Orang Asli dari Semenanjung Malaysia dan untuk BangsaBangsa Bumiputera dari Sabah, Sarawak dan Brunei. BAHAN DAN CARA Seperti oleh Tan (978). HASILHASIL Data yang telah dikumpulkan disenaraikan dalam jadualjadual berikut. Jadual menunjukkan data untuk beberapa kumpulan darah dan juga untuk sekresi bahanbahan kumpulan darah ABH ke dalam air liur (taraf sekresi). Jadual 2 menunjukkan tandatanda genetik biokimia dari serum, Jadual 3 dari air liur dan Jadual 4 dari eritrosit. Jadual 5 ialah suatu senarai tandatanda genetik biokimia yang tidak menunjukkan varian dalam bangsabangsa Malaysia di bawah perhatian dan Jadual 6 menunjukkan tandatanda genetik biokimia yang hanya telah diuji di dalam orang Kadazan. JADUAL Kumpulan darah. (a) ABO Bangsa Tempat Bil. IA Frekuensi gen IB IO Rujukan Melayu China India India Utara India Selatan Negrito Senoi Melayu Asli Orang Asli Bidayuh Iban Melanau Kadazan Kedayan Kuala Lumpur Malaya Singapura Brunei Kuala Lumpur Malaya Singapura Kuala Lumpur Malaya Singapura Malaya Sarawak Sabah Brunei 4, ,46, , , , 6, ,, Duraisamy dan Amarasingam, 97 Lum dan Amarasingam, 968 Chan, 962 Allen dan ScottMacGregor, 947 Graydon ail, 952 Duraisamy dan Amarasingam, 97 Lum dan Amarasingam, 968 Simmons dll., 956 Chan, 962 Allen dan ScottMacGregor, 947 Yeoh, 96 Duraisamy dan Amarasingam, 97 Lum dan Amarasingam, 97 Chan, 962 Allen dan ScottMacGregor, 947 Allen dan ScottMacGregor, 947 LieInjo, 976 LieInjo, 976 LieInjo, 976 Simmons dll., 95 Graydon dll., 952 Polunin dan Sneath, 953 Graydon dll., 952 Graydon dll., 952 Ride, 932 Graydon dll.,

31 TANDATANDA GENETIK KUMPULANKUMPULAN DARAH MALAYSIA BRUNEI SINGAPURA JADUAL (b) MN. Frekuensi gen Bangsa Tempat Bil. M N Rujukan Melayu Kuala Lumpur Singapura LieInjo, 976 GibsonHill, 953 Brunei Graydon dll., 952 China Kuala Lumpur Malaya Steinberg dll., 96 Simmons dll., 95 India Negrito Senoi Melayu Asli Bidayuh Kuala Lumpur Malaya Sarawak Steinberg dll., 96 Polunin dan Sneath, 953 Polunin dan Sneath, 953 Polunin dan Sneath, 953 Polunin dan Sneath, 953 Graydon dll., 952 Iban Sweetman dll., 95 Graydon dll., 952 Melanau Kedayan Brunei Graydon dll., 952 Graydon dll., 952 JADUAL (c) Rhesus. Frekuensi gen Bangsa Melayu Tempat Singapura Bil. 42 CDE (Rz) CDe (Ri) cde (R2) cde (Ro).8 cde (r) Rujukan GibsonHill, 953 Brunei Polunin dan Sneath, 953 China Malaya Simmons dll., 95 Kuala Lumpur Steinberg dll., 96 India Negrito Malaya Steinberg dll., 96 Polunin dan Sneath, 953 Senoi Polunin dan Sneath, 953 Melayu Asli 6 _ Polunin dan Sneath, 953 Bidayuh Sarawak Polunin dan Sneath, 953 Iban Graydon dll., 952 Graydon dll., 952 Melanau Graydon dll., 952 Kedayan Brunei Graydon dll.,

32 S. G. TAN JADUAL (d) P Frekuensi gen Bangsa Tempat Bil. Pi P2 p Rujukan Negrito Malaya Polunin dan Sneath, 953 Senoi Polunin dan Sneath, 953 Bidayuh Sarawak Polunin dan Sneath, 953 Melanau.5.5 Graydon dll., 952 Kedayan Brunei Graydon dll., 952 JADUAL (e) Lewis (Le a ) Fenotaip (Nombor dan %) Bangsa Tempat Bil. Le(a + ) Le(a) Rujukan Melayu Singapura 93 23(24.7%) 7(75.3%) Saha dll., 973 China 24 33(5.4%) 8(84.6%) Saha dll., 973 India 8 25(2.2%) 73(78.8%) Saha dll., 973 Melanau Sarawak 4(4.%) 86(86.%) Graydon dll., 952 Kedayan Brunei 4 9(8.3%) 85(8.7%) Graydon dll., 952 JADUAL (f) Xg* Fenotaip (Bilangan dan n Lelaki sahaja o) Bangsa Tempat Bil. Xg(a + ) Xg(a ) Rujukan Melayu China Singapura (57.%) 4(47.%) 2(42.9%) 46(52.9%) Saha dll., 973 Saha dll., (45.7%) 9(54.3%) Boon dll., 964 India 85 48(56.5^,,) 37(43.5%) Saha dll., 973 (g) Taraf JADUAL Sekresi (Secretoi Status) Frekuensi gen Bangsa Tempat Bil. Se se Rujukan Negrito Malaya Polunin dan Sneath, 953 Senoi Polunin dan Sneath, 953 Melayu Asli Bidayuh Kedayan Sarawak Brunei Polunin dan Sneath, 953 Polunin dan Sneath, 953 Graydondll.,

33 TANDATANDA GENETIK KUMPULANKUMPULAN DARAH MALAYSIA BRUNEI SINGAPURA JADUAL 2 Tandatanda genetik biokimia dari serum. (a) Haptoglobin (Hp) Bangsa Tempat, Bil. Frekuensi gen Hpl Hp2 Rujukan Negrito Malaya.5.95 LieInjo, 976 Senoi LieInjo, 976 Melayu Asli LieInjo, 976 Orang Asli LieInjo dll., 967a Bidayuh Sarawak Ganesan dll., 975a Iban Ganesan dll., 975a Kadazan Sabah Tan dll., 978 (b) JADUAL 2 Transferrin ( Tf) Frekuensi gen Bangsa Tempat Bil. TfC TfD Rujukan Orang Asli Malaya LieInjo dll., 967b Bidayuh Sarawak Ganesan dll., 975a Iban Ganesan dll., 975a Kadazan Sabah Tan dll., 978 (c) JADUAL 2 Albumin (Alb) Bangsa Tempat Bil. Jenis dan nombor varianvarian nadir Rujukan Bidayuh Sarawak 283 Ganesan dll., 975a Iban 88 N + Medan () Ganesan dll., 975a Orang Asli Malaya 65 N + Gombak (2) LieInjo dll., 97 Bangsa (i) SapA Senoi Melayu Asli (ii) SapB Senoi Melayu Asli Tempat Malaya Malaya JADUAL 3 Tandatanda genetik biokimia dari air liur. (a) Fosfatase acid air liur. Bil. Frekuensi gen Rujukan A B Al. B*.. A B.9 _ Tan dan Teng, 978 Tan dan Teng, 978 Tan dan Teng, 978 Tan dan Teng, 978

34 S. G. TAN JADUAL 3 (b) Superoksid dismutase B (SOD B) Frekuensi gen Tempat Bil. SOD Bl SOD B2 Rujukan Senoi Malaya Tan dan Teng, 978 Melavu Asli 32. Tan dan Teng, 978 JADUAL 4 Tandatanda genetik biokimia dari eritrosit. (a) Glukos6fosfat dehidrogenase (G6PD): Kekurangan aktiviti enzim didalam eritrosit. Lelaki sahaja Bangsa Tempat Bil. Nombor dengan kekurangan (t () dengan kekurangan Rujukan Melavu Senoi Melavu Asli Orang Asli Bidayuh I ban Kadazan Bajau Murut Bisaya Brunei Sarawak Sabah Malaya Sarawak Sabah LieInjo dll., 964 LieInjo du., 964 LieInjo dll., 964 LieInjo dan Chin, 964 LieInjo dan Chin, 964 LieInjo dan Chin, 964 LieInjo dan Ti, 964 LieInjo dll., 964 Ganesan dll., 975b LieInjo dll., 964 Ganesan dll., 975b LieInjo dll., 964 LieInjo dll., 964 LieInjo dll., 964 LieInjo dll., 964 JADUAL 4 (b) Fosfoglukomutase I (PGMi) Frekuensi gen Bangsa Tempat Bil. PGMj PGM? Rujukan Negrito Malaya LieInjo, 976 Senoi LieInjo, 976 Melavu Asli LieInjo, 976 Orang Asli Welch dll., 972 Bidayuh Sarawak Ganesan dll., 976 Iban Ganesan dll., 976 Kadazan Sabah Tan dll., 978 9

35 TANDATANDA GENETIK KUMPULANKUMPULAN DARAH MALAYSIA BRUNEI SINGAPURA JADUAL 4 (c) Adenosin deaminase (ADA) Bangsa Tempat Bil. Senoi Malaya ^22 Melayu Asli 64 Bidayuh Sarawak 27 I ban 88 Kadazan Sabah 283 Frekuensi gen ADA* ADA2 Rujukan Welch dll., 978,75 Welch ail., Ganesan ail., Ganesan ail,, Tan dll., 978 Bangsa Tempat Bil. (d) JADUAL 4 Adenilat kinase (AKi) Frekuensi gen AK Rujukan Senoi Melayu Asli Malaya LieInjo, 976 LieInjo, 976 Orang Asli Welch dll., 97 Bidayuh Sarawak 27. Ganesan dll., 976 I ban Ganesan dll., 976 Kadazan Sabah 25. _ Tan dll., 978 JADUAL 4 (e) 6F*osfoglukonat dehidrogenase (6PGD). Bangsa Tempat Bil. Negrito Malaya Senoi Melayu Asli Orang Asli 253 Bidayuh Sarawak 32 Iban 27 Kadazan Sabah 284 Frekuensi gen PGDA PGDC Rujukan LieInjo, 976 LieInjo, 976 LieInjo, 976 LieInjo dan W'elch, 972 Ganesan dll., 976 Ganesan dll., 976 Tan dll., 978 JADUAL 4 (f) Peptidase B (PEP B) Bangsa Tempat Bil. PEP Bi Frekuensi gen PEP B6 PEP B2 Rujukan Senoi Malaya Welch, 973 Melayu Asli Welch, 973 Kadazan Sabah _ Tan dll.,

36 S. G. TAN JADUAL 4 (*) Hemoglobin (Hb) Jenis dan nombor Bangsa Tempat Bil. varianvarian naui^ Rujukan Melayu Sarawak 66 AE(5) Vella dan Tavaria, 96 Senoi Malaya 73 AE(44)EE(5) LieInjo dan Chin, AE(3)EE(22) LieInjo dll., 973 Orang Asli B 2 (l), AE(32) EE(43) AE(278)EE(45) LieInjo, 97 Welch dll., AE(425)EE(7) LieInjo dll., 972 Melayu Asli 75 AE(9) LieInjo dan Chin, AE(7)ACoSp(9) LieInjo dll., 973 Bidayuh Sarawak 285 ACoSp(l) Ganesan dll., 975b AE() Colbourne dll., Vella dan Tavaria, 96 Iban 24 ACoSp(2) Ganesan dll., 975b 85 Colbourne dll., Vella dan Tavaria, 96 Murut Sabah 53 AE(3) Vella dan Tavaria, 96 JADUAL 5 Tandatanda genetik biokimia yang tidak menunjukkan varian pada bangsabangsa Malaysia yang telah diuji. Tandatanda genetik dan sumber Amilase (AMY2) serum Laktat dehidrogenase (LDH) eritrosit Karbonik anhidrase (CA) eritrosit Malat dehidrogenase (MDH) eritrosit Superoksid dismutase A (SOD A) air liur Amilase (AMYl) air liur Fosfoheksos isomerase (PHI) eritrosit Giutamat oksaloasetik transaminase (sgot) eritrosit Superoksid dismutase A (SOD A) eritrosit Bangsa dan bil. yang telah diuji Kadazan (254) Bajau (6) Bidayuh (285) Iban (22) Bidayuh (283) Iban (29) Orang Asli (534) Orang Asli (32) Melayu Asli (32) Senoi (23) Senoi (5) Melayu Asli () Orang Asli (2) Bidayuh (285) Iban (28) 93 Kadazan (266) Kadazan (233) Rujukan Teng dll., 978 Ganesan dll., 976 Ganesan dll., 976 Welch dll., 972 Welch dll., 97 Tan dan Teng, 978 Tan dan Teng, 978 LieInjo dan W T elch, 972 Ganesan dll., 976 Tan dll., 978 Tan dll., 978

37 TANDATANDA GENETIK KUMPULANKUMPULAN DARAH MALAYSIA BRUNEI SINGAPURA JADUAL 6 Tandatanda genetik biokimia darah yang telah diuji dalam Orang Kadazan (Tan dll., 978). Tandatanda genetik Glioksalase I (GLO) Komponen kumpulan khas (Gc) Esterase D (ESD) Bil Frekuensi gen GLOl =.3; GLO2 =.7 Gcll =.868; Gc2 =.32 ESDI =.472; ESD2 =.528 Fosfatase asid (ACPi) 264 P2 =>.369; P b «=.63 Glutamik piruvik transaminase (sgpt) Uridine monotosfiit kinase (UMPK) Peptidase D (PEP D) C GPTi «.265; GPT2 =.735 UMPKl.959; UMPR2.4 PEP Di.994; PEP D3 =.6 Insiden C5 2.7% PERBINCANGAN Kekurangan data genetik, terutama sekali untuk tandatanda genetik biokimia dalam bangsabangsa Bumiputera Sabah, Sarawak dan Brunei dan juga dalam Orang Asli Semenanjung Malaysia menjadi jelas jika kita membandingkan data genetik yang dipersembahkan di sini dengan data yang telah disusun untuk orangorang Melayu, China dan India dari Semenanjung Malaysia (Tan, 978). Tidak terdapat, atau hampirhampir tidak terdapat, apaapa data genetik untuk beberapa bangsa di Sabah, Sarawak dan Brunei seperti Murut, Bajau, Melanau, Kayan, Kajang, Kenyah, Kelabit, Penan dan Punan. Untuk Iban, Bidayuh dan Orang Asli, sungguh pun data ada untuk beberapa kumpulan darah dan tandatanda genetik biokimia, tetapi data tidak ada untuk beberapa tanda genetik biokimia baru yang sangat polimorfik dan oleh yang demikian amat berguna terhadap pengajian antropologi dan genetik seperti glioksalase I, esterase D dan glutamik piruvik transaminase. Oleh itu, banyak lagi penyelidikan harus dibuat ke atas bangsabangsa tersebut sebelum suatu ketahaman tentang strukturstruktur genetik mereka boleh dicapai. Mungkin penyelidikanpenyelidikan seperti ini boleh dibuat di bawah anjuran Universiti Asean apabila ditubuhkan kelak. Penubuhan Universiti ini telah dicadangkan tidak lama dahulu dan kampusnya mungkin terletak di Labuan, Sabah. Orang Melayu, Orang Asli dan Bangsa Bangsa Bumiputera Sabah, Sarawak dan Brunei lebih menarik dari segi antropologi bila dibandingkan dengan orangorang China dan India di Malaysia, sebab mereka adalah bangsabangsa Asia Tenggara. Sebaliknya orangorang China 94 dan India di Malaysia adalah kumpulankumpulan imigran. Tanah besar Asia Tenggara ialah suatu semenanjung timur dari negeri India dan selatan dari negeri China; kepulauan Asia Tenggara termasuk pulaupulau Indonesia, Filipina dan Taiwan. Pulaupulau ini membentuk suatu jambatan di antara benua Asia dan benua Australia dan mereka memisahkan lautan Hindi dan Pasifik. Daerah ini mempunyai lebih ramai bangsa dan kebudayaan yang berbeza bila dibandingkan dengan Iainlain daerah yang mempunyai saiz yang sama. Misalnya, lebih dari 3 bangsa yang berlainan boleh dijumpai ditanah besar Asia Tenggara sahaja dan,2 nama telah digunakan untuk merujuk terhadap mereka (Provencher, 975). Bila kepulauan Asia Tenggara juga diambil perhatian, bilangan bangsabangsa ini melambung naik (Lebar, 972, 975). Oleh yang demikian penyelidikanpenyelidikan antropologi fizikal dan genetik populasi mungkin sangat berfaedah dan memuaskan di daerah ini. Malaysia, yang terletak di pertengahan Asia Tenggara dan yang mengandungi di dalam wilayahnya keduadua tanah besar Asia Tenggara (Semenanjung Malaysia) dan kepulauan Asia Tenggara (negerinegeri Sabah dan Sarawak), ialah tempat yang sesuai sekali untuk membuat penyelidikan antropologi dan genetik manusia sebab di antara pendudukpenduduknya boleh dijumpai ramai bangsabangsa asal keduadua tanah besar Asia Tenggara dan kepulauan Asia Tenggara. Saya berhar?.p kertas ini dan juga kertas saya yang dahulu (Tan, 978) akan lebih menggalakkan lagi pengk?jian genetik dan antropologi dalam bangsabangsa yang menjadikan populasi Malaysia sekarang, terutama sekali penyelidikan yang dibuat oleh anak negeri kita sendiri.

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39 TANDATANDA GENETIK KUMPULANKUMPUL AN DARAH MALAYSIA BRUNEI SINGAPURA STEINBERG, A.G., LAI, L.Y.C., VOS, G.H., BHAGWAN SINGH, R., LIM, T.W. (96): Genetic and population studies of the blood types and serum factors among Indians and Chinese from Malaya. Am. J. Hum. Genet. 3: SWEETMAN, K.E.D., IKIN, E.W., MOURANT, A.E. (95): Dalam: "The distribution of human blood groups and other polymorphisms''. oleh A.E. Mourant, A.C Kopec, K. Domaniewska Sobczak. Edisi kedua. Oxford University Press. London TAN, S.G. (978): Tandatanda genetik biokimia dalam tiga bangsa terbesar disemenanjung Malaysia dan Smgapura: Satu penyusunan data. Pertanika. : TAN, S.G. dan TENG, Y.S. (978): Saliva acid phosphatases and amylase in Senoi and Aboriginal Malays and superoxide dismutase in the various racial groups of Peninsular Malaysia. Jpn< J. Hum. Genet. (Dalam pencetakan). TAN, S.G., TENG, Y.S., GANESAN, J., LAU, K.Y., LIE INJO, L E. (978): Biochemical genetic markers in the Kadazans of Sabah, Malaysia. Hum. Genet. (Dihantarkan). TENG, Y.S., TAN, S.G., LOPEZ, C.G., NG, T., LIE INJO, L.E. (978): Genetic markers in Malaysians: Variants of soluble and mitochondrial glutamic oxaloacetic transaminase and salivary and pancreatic amylase. Phosphoglucomutase III and saliva esterase polymorphisms. Hum. Genet. 4: VELLA, F. dan TAVARIA, D. (96): Haemoglobin variants in Sarawak and North Borneo. Nature. 9: WELCH, Q.B. (973): Peptidase B variants among the Semai, Temuan, Semelai and Jakun groups of West Malaysian Orang Asli. Hum. Hered. 23: 482^86. WELCH, Q.B., LIEINJO, L.E., BOLTON, J.M. (97): Adenylate kinase and malate dehydrogenase in four Malaysian racial groups. Humangenetik 4:663. ' WELCH, Q.B, LIEINJO, L.E., BOLTON, J.M. (972): Phosphoglucomutase and carbonic anhydrase in West Malaysian Aborigines. Hum. Hered. 22: WELCH, Q.B., LEE, C.S., THANGAVELU, S., LIEINJO, L.E. (978): Adenosine deaminase polymorphism among the Semai, Temuan, Semelai and Jakun groups of West Malaysian Orang Asli. Hum. Hered. 28:6265. WILLIAMHUNT, P.D.R. (952): An introduction to the Malayan Aborigines. Government Press. Kuala Lumpur. (Received 26 July 978) 96

40 Pertanika (2), 974 (978) Effects of Selection for High LitterWeight on Reproductive Performance in Mice K. BAHARIN and R. G. BEILHARZ 2 Faculty of Veterinary Medicine and Animal Science, Universiti Pertanian Malaysia Key words: Reproductive Performance, Mice, Litterweight, Selection, Inbreeding. RINGKASAN Prestasi pembiakan dari dua kumpulan tikus yang dipilih kerana berat litter ketika berumur sembilan minggu dan satu kumpulan kawalan telah dibandingkan hasil dari data yang diperolehi selepas 5 keturunan pembiakan. Dua puluh pasang baka dibiakkan bagi kumpulan kawalan dan sepuluh pasang untuk kumpulankumpulan yang dijalankan pemilihan. Pada setiap keturunan baka hanya dikahwinkan sekali. Dari kirakira angkali pembiakan sedarah (inbreeding) menunjukkan kadar kenaikan angkali pembiakan sedarah yang tinggi di kalangan kumpulan yang dijalankan pemilihan. Keputusan menunjukkan kurangnya kesan (response) yang dihasilkan dari pemilihan. Sebaliknya prestasi pembiakan telah merosot bersamaan dengan kenaikan angkali inbreeding di kalangan kumpulan yang dibuat pemilihan walaupun pertalian di antara kedua faktor ini tidak linear. Penambahan dari segi kenaikan berat badan dilihat pada peringkat awal. Akan tetapi apabila angkali inbreeding melebihi 4% sifat ini juga turut merosot. SUMMARY The reproductive performances of two lines of mice selected for high litterweight at nine weeks of age and a control line were compared based on data obtained after 5 generations of breeding. Twenty pairs of parents were bred per generation in the control line and ten pairs in each of the selected lines. A t each generation the parents were mated only once. Calculation of inbreeding coefficients from pedigrees indicated a rapid rise in inbreeding coefficient in the selected lines. There was little response to selection for high litterweight and in fact there was a general decline in reproductive performance associated with high levels of inbreeding in the selected lines although the relationship was not linear. Some initial response in body weight gain was observed but even this trait showed a decline in later generations when inbreeding coefficient exceeded 4%. INTRODUCTION Reproduction in a female mammal may be defined as its ability to produce offspring and raise them till they become independent of the parents. In mice reproduction is measured usually in terms of littersize or total litterweight. As in other traits, reproduction may be subjected to selection, both natural and artificial. Natural selection is always acting to maximise fitness. It has been suggested that because the capability of animals to reproduce has been under natural selection throughout the evolutionary history of the species, little additive genetic variation is left and as a result we may Part of Ph. D. Thesis submitted to the University of Melborne. 2 School of Agriculture and Forestry, University of Melbourne expect it to be rather difficult to make marked genetic improvement (Robertson, 963; Politiek, 965; Karg, 969). However, there is still genetic variation for reproductive traits as shown by successful selection experiments in Merino Sheep (Turner and Young, 969). This study is aimed at analysing the direct and indirect responses to selection for high total litterweight at nine weeks of age in laboratory mice. Traits analysed in this study include littersize at birth, total litterweight at three weeks and at nine weeks of age, average body weight at three and nine weeks of age, and mortality of progeny between birth to three weeks and from three weeks to nine weeks of age.

41 MATERIALS AND METHODS The analysis was based on the records of performance at mating and subsequent growth of the litters prcduced by mice in the selected population (line 6A and line 6B) and the control population (line ). The origin of the stock and the management practices have been documented by Beilharz (97, 972). The original mice stock came from the Division of Animal Health, C.S.I.R.O., Parkville, Melbourne, where a population of mice was propagated without conscious selection, by at least 4 males and 6 females per generation. This population should not have been inbred to any great extent. Of the 5 mice obtained from C.S.I.R.O., 2 males and 79 females were mated to found the mice colony at the School of Agriculture, University of Melbourne. These mice were designated as generation and their progeny as generation. The mice in generation were divided to two populations at random, 37 litters into each of populations A and B. In both populations all litters were reduced to 6 at birth by random removal of the excess young. Population A was fully fed throughout. Mice from population B were fed a restricted amount (about 8% of the estimated full intake) between 3 and 9 weeks of age. At other times feeding in population B was ad lib. as for population A. Fifty pairs of mice were mated in each population to produce generations 2 and 3. In generation 4, and in all subsequent generations, all mice were fed ad lib. A total of 582 mice were identified in generation 4 and they were all treated as one population. One control line (line ) and 2 selected lines (including lines 6A and 6B) were separated from this population to produce generation 5. Line is an unselected population propagated by 2 pairs of mice each generation. Each pair was called a family (numbered from to 2). One son and one daughter are chosen, at random from each family for breeding. If a family (say family n) fails to produce a son or daughter, a substitute is obtained from the family n+9 or n+. While daughters remained in the family of their parents the sons were moved systematically to other families. In this way if all families leave a son and a daughter, the possibility of inbreeding is avoided for five generations Lines 6A and 6B were selected lines where males and females were selected on high total weight of licter at 9 weeks of age. The only difference between lines 6A and 6B was that in line 6B mating of full sibs was avoided whenever K. BAHARIN AND R. G. BEILHARZ 98 possible. However, because of the family selection employed it was not always possible to avoid mating full sibs. Ten pairs of mice were mated at each generation. Variation in sex ratio was allowed for as follows: the sex difference (malefemale) was calculated from the average weight of all male and female mice from line, line 6A and line 6B. This average difference was then added to the weight of each female. The following records were collected from the selected and control lines on a routine basis: litter size (number of live young born), weight at 3 weeks (weaning weight), weight at 6 weeks, weight at 9 weeks, sex ratio in the litter, matings (and hence, pedigree), and mortality. RESULTS The coefficient of inbreeding for each individual at each generation was estimated using the path coefficient technique (Wright, 922). Table shows the mean changes in the coefficient of inbreeding over the generations. In line, inbreeding started to increase at generation 8 and 9 but dropped at generation, increased steadily from generation to 4 and showed a second drop in generation 5 before the next wave of increase from generation 6 to 8. In lines 6A and 6B, inbreeding coefficient increased rapidly from generation 6 onwards. The decrease in inbreeding coefficient observed in Line 6A from generation 6 to generation 7 was due to larger number of families selected to provide sufficient individuals to become parents for mice in generation 7 compared to that for generation 6. Table 2 shows the percentage of successful mating in each generation. Mice in Line continued to produce offsprings at each generation although a few families (maximum observed for any generation was 3 families) failed to produce any offsprings. In Line 6A, successful matings were maintained at 9 % up to generation 8 after which there was a gradual decline and at generation 3 no offspring was produced and the line became extinct. In Line 6B, deterioration of fertility was observed at generation 4 when the percentage of successful mating dropped to 7%. Although there was a recovery at the next generation, the fertility continued to drop at generation 6 to 8 and at generation 9 no offspring was produced and the line became extinct. Table 3 shows the intensity of selection (i) estimated as selection differential divided by standard deviation (Falconer, 96) applied at

42 SELECTION FOR HIGH LITTERWEIGHT ON REPRODUCTIVE PERFORMANCE IN MICE TABLE Changes in coefficient of inbreeding Average % of inbreeding within each line Generation Average for all litters Average for selected mice Line Line 6A Line 6B Line Line 6A Line 6B TABLE 2 Percentage of successful mating in each generation Generation Line % Line 6A % Line 6B % each generation for the selected trait (which in this case is the litterweight at nine weeks of age) and its response. The results indicate that the only period when there was some indication of response to selection was in the earlier generations (generation 8 to ) when there was a tendency for both Line 6A and Line 6B to increase in their total litter weight. The weight subsequently dropped below to that of Line. Among the other traits, litter size at birth among the selected lines did not differ very much from that of the control line until just before the extinction of the selected lines. The weight of litters at weaning showed a similar trend to that of the total litter weight at 9 weeks of age. Mortality rate of mice was very high among the selected lines at the later generation when the coefficient of inbreeding was high but the relationship of the two factors was not linear.

43 K. BAHARIN AND R. G. BEILHARZ TABLE 3 Mean adjusted total litterweight at nine weeks in each generation in the unselected and selected lines, and the intensity of selection applied Line Line 6A Line 6B Generation (of parents) (control) Total Litter wt. (grams) Selection intensity (i) Total litterweight (grams) Selection intensity (i) Total litterweight (grams) ± ± ± ± ± : : ± ± ;: ± ± t ± ± ± ± ± ± ± b ± ± ± ± ± t: ± ± ± ± : : TABLE 4 Average littersize (live and dead) at birth in each generation Generation (of parents) Line Line 6A Line 6B TABLE 5 Adjusted total litterweight at three weeks in each generation in the various lines Generation (of parents) Line Line 6A Line 6B g g g

44 SELECTION FOR HIGH LITTERWEIGHT ON REPRODUCTIVE PERFORMANCE IN MICE TABLE 6 Percentage mortality between birth and three weeks, and between three to nine weeks of age in each generation Generation (of parents) Line % Mortality, birth to 3 weeks % Mortality, 3 to 9 weeks Line 6A Line 6B Line Line 6A Line 6B The mean body weight of the selected progeny generation when the coefficient of inbreeding at three weeks of age showed a decline after exceeded 25% in both Lines 6A and 6B. Generation TABLE 7 Mean bodyweight at three weeks of age in each generation among males and females selected for breeding in the various lines Males.lg Line Females.5g Males 3.4g Line 6A Females 3.7g Males 3.3g Line 6B Females 3.lg

45 K. BAHARN AND R. G. BELHARZ The mean bodyweight at nine weeks of age tended to decline after generation 3 in line 6B when the inbreeding coefficient exceeded 45%. There was no marked decline in bodyweight at nine weeks of age in line 6A. DISCUSSION The differences in the response of the two selected lines may in part be due to some differences in the genetic constitution of the two lines although every effort had been made to give similar treatment to both lines. As described previously the control line and the selected lino were derived from the progeny of two populations A and B where some differences in feeding were imposed at generations 2 and 3. At generation 4, all the progeny from both population were mixed and treated as one population. From this base population, the various lines were developed. Because of the random distribution of mice into the various lines it was assumed that all the lines would have a similar genetic constitution. This assumption may not have been valid particularly when the size of each line was rather small (only families for the selected lines) as this could result in significant variation in gene frequency between lines due to the effect of sampling (Falconer, 96). The assumption that mice of generation 4 were unrelated is also not correct. The effect of the assumption is to make the coefficient of inbreeding in the early generations of Table lower than it actually is. However, increase in the inbreeding coefficient after generation 5 is no longer affected by this assumption. To obtain a reasonable response from selection the intensity of selection must be high and this requires heavy culling of animals at each generation. This same procedure will also lead to reduction in population size resulting in a rapid rise in inbreeding coefficient with its consequent depressing effects on performance. The two opposing factors acted jointly in line 6A and line 6B and it was not possible to estimate the separate effects of each. Looking at the changes in the level of performance at each generation it may be inferred that the only trait to indicate some response to selection was the growth rate of the individuals in the litter when the rate of inbreeding was still low. Similar results were Generation TABLE 8 Mean bodyweight at nine weeks of age in each generation among selected for breeding in the various lines Males 3.9g , Line Females 28.7g Males 35.2g Line 6A Females 29.8g males and females Males 36.Ig Line 6B Females 3.g

46 SELECTION FOR HIGH LITTERWEIGHT ON REPRODUCTIVE PERFORMANCE IN MICE reported in Iowa with experiments in pigs where continuous selection and inbreeding had been applied (Bereskin et al, 969, 97). Selection was shown to be effective and inbreeding less depressing on growth while littersize showed an erratic but generally downward trend. Most experiments on selection for high rate of reprcduction in various species of domestic mammals have been shown to be unsuccessful except in sheep (Revelle and Robinson, 973). Dalton and Bywater (963) obtained no response to selection for littersize and litterweight at weaning age after 4 generations of selection in mice maintained at high and low levels of feeding. Some workers, however, reported some indirect response through selection for bodyweight (Fowler and Edward, 96; Rahnefdd et al, 968). Bradford (97, 97), however, obtained no response in littersize either through direct selection for the trait or through indirect selection for weight gain between 2 and 42 days of age but there was a response to selection for low litter size. The general decline in reproductive performance among the selected lines may be related to the depressing effect of inbreeding but it could also be due to the fact that the animal's overall fitness has been compromised by conflicting selection pressures as suggested by Goddard and Beilharz (977), according to whom there are certain optimum values and combinations for the various components that constitute total fitness. Selection for one component of fitness may have caused a decline in the total fitness. This has been demonstrated in sheep where there has been a definite cost associated with selection for twins among Corriedale sheep (Baharin and Beilharz, 977a). Baharin and Beilharz (977b) showed that in pigs there was a high percentage of mortality at very large littersizes and that little is to be gained by increasing the total litter size above 3 piglets when artificial rearing is practised. CONCLUSIONS 3 Selection for high litterweight at nine weeks of age in mice failed to show any positive response. In fact, it showed a depression in performance compared to the performance in the control line. This could be due to inbreeding depression or breakdown in total fitness due to conflicting selection pressures. Fertility as measured by mortality rates of progeny between birth and three weeks of age, and between three and nine weeks of age, was unaffected by selection and depressed at high levels of inbreeding. Bodyweight showed some initial response at the beginning of the selection programme, but at very high level of inbreeding, this trait too showed depression. ACKNOWLEDGEMENTS We are grateful to Miss Susan Clarke and Mr. B. Coulson for the collection of the data. The work was supported by the Australian Meat Research Committee. REFERENCES BAHARIN, K. and BEILHARZ, R.G. (977b): A Comparison of the performance of single and twin born corriedale ewes and lambs. Aust. J. exp. Agric Anim. Hush. 7: BAHARIN, K. and BEILHARZ, R.G. (977a): An analysis of reproductive performance in pigs based on records of performance of the boar. Aust. J. Exp Agric. Anim. Husb. 7: BEILHARZ, R.G. (97): Evaluation of continual crossbreeding with inbred lines of mice. Proc Aust. Soc. Anim. Prod. 8: BEILHARZ, R.G. (972): "Studies on Mouse Breeding." Occasional Paper No.. School of Agriculture, University of Melbourne. BERESKIN, B.B., SHELBY, C.E. and HAZEL, L.N. (969): Monte Carlo studies of selection and inbreeding I. Genetic and phenotypic trends. J. Anim. Sci. 29: BERESKIN, B.B., SHELBY, C.E. and HAZEL, L.N. (97): Monte Carlo studies of selection and inbreeding. II. Inbreeding coefficient. J. Anim. Sci. 3: BRADFORD, G.E. (97): Reproduction in mice selected for rapid growth. J f. Anim Set. 3: 6. BRADFORD, G.E. (97): Growth and reproduction in mice selected for rapid bodyweight gain. Genetics. 69: DALTON, D.C. and BYWATER, T.L. (963): The effect of selection for litter size and litterweight at weaning in mice maintained on two diets. Anim. Prod. 5: FALCONER, D.S. (96): ''Introduction to Quantitative Genetics." Edinburgh. Oliver add Boyd 365 pp. FOWLER, R.E. and EDWARDS, R.G. (96): The fertility of mice selected for large or small body size. Genet. Res. : GODDARD, M.E. and BEILHARZ, R.G. (977): Natural selection and animal breeding. Proc. 3rd. hit. Cong. SABRAO. Canberra, pp. 4(9). KARG, H. (969): Opportunities for improving reproduction as a mean to increasing efficiency of animal production. Proc. 2nd World Conf. Anim. Prod., Maryland, pp. 29. POLITIEK, R.A. (965): Fertility as a breeding problem. World Rev. Anim. Prod. : 5964.

47 K. BAHARIN AND R. G. BEILHARZ RAHNEFELD, G.W., COMSTOCK, R.E., SINGH, M. and ROBERTSON, H. (963): Inbreeding in artificial selection NAPUKET,S.R.(966): Genetic correlation between programmes. Genet. Res. 2: growth rate and litter size in mice. Genetics. 54: XT, _., /4rt lt^ TURNER, H.N. and YOUNG, S.S.Y. (969): "Quantitative Genetics in Sheep Breeding." Melbourne. MacMillan. 332 pp. REVELLE, T.J. and ROBINSON, O.W. (973): An explanation for the low heritability of littersize in WRIGHT, S. (922): Coefficient of inbreeding and swine. J. Anim. Set. 37: relationship. Amer. Nat. 56: (Received 9 August 978) 4

48 Pertanika (2), 5 (978) Tax Burden on Rubber, Coconut and Pineapple Smallholders in Johore MOHAMMED ARIFF BIN HUSSEIN Faculty of Resource Economics and Agribusiness, Universiti Pertanian Key words: Tax burden, Smallholders RINGKASAN Malaysia Kajian ini meneliti samada cukaicukai yang dikenakan keatas pekebunpekebun kecil getah, kelapa dan nenas di tahun 975 menjadi sebab utama kerendahan pendapatan mereka. Hasil kajian menunjukkan yang beban cukai keatas pekebunpekebun kecil adalah besar, terutama sekali kepada pekebunpekebun kecil get ah. Pekebunpekebun getah biasanya membayar cukai sebanyak seperempat hingga sepertiga daripada pendapatan mereka. Bagaimana pun kemungkinan menambah pendapatan dengan car a mengubah struktur cukai adalah terbatas. Perubahanperubahan begitu hendaklah disertakan dengan programprogram yang akan meninggikan day a pengeluaran dan luas kebun. SUMMARY The study examines whether taxes levied on rubber, coconut, and pineapple smallholders in 975 are a major cause for smallholder low income. Results of the study show that tax burden on the smallholder is substantial, particularly that on rubber smallholders. The average rubber smallholder pays about onefourth to onethird of his income in taxes. However, the scope for increasing income through changes in the tax structure is limited. Such changes should be coupled with programmes aimed at increasing farm productivity and size of holding. INTRODUCTION In Peninsular Malaysia 2 overall equity as measured by the Gini Concentration ratio, increased from.42 in 957 to.52 in 97. The low income groups also experienced a decline in their absolute income levels. The mean income of the lowest 4 percent of households decreased from $86 per month in 958 to $75 in 97 (Snodgrass, 975). Largely in response to this problem, the government has embarked on programmes aimed at increasing the income level of the poor, particularly those in the agricultural sector. These programmes include land development schemes and in situ development projects such as the provision of irrigation and drainage facilities, replanting of agricultural crops with new high yielding varieties, and improvements in the marketing outlets for farm products. A neglected aspect of the problem is the effect of taxes on the poor in the agricultural sector. Studies on tax incidence in the country show that the tax system is highly regressive at the lower end of the income scale. The regressivity was attributed mainly to the export duty on rubber through its effect on smallholders (McLure, 972), and to import duties and excise taxes (Snodgrass, 975). Certain taxes, notably landbased taxes, however, were not considered in both of the studies. The inclusion of these taxes could have added to the regressivity of the tax incidence for the low income groups. Conceivably one of the causes for low income in the agricultural sector is the high tax burden on the poor in that sector. The objective of this study is to estimate the tax burden on rubber, coconut and pineapple smallholders of less than 5 acres in Johore in their capacity as producers and purchasers of production inputs. 3 CONCEPTUAL FRAMEWORK Tax incidence is defined as the dollar burden of a tax distributed among different economic Part of Ph.D. Thesis submitted to Pennsylvania State University, Unless otherwise stated subsequent references to Malaysia in this paper refers only to Peninsular Malaysia. 3 The base year of this study is 975. All estimates of income were adjusted to 975 and changes in the relevant taxes after 975 were not included in the study. Smallholders in this study refer only to the traditional scattered smallholders. 5

49 units. The burden of a tax is measured as a ratio of the amount of tax paid and income. Three measures of income were used gross income, income net of all costs except family labour, and income net of variable costs. The method used in analysing tax shifting and incidence was one of deductive partial equilibrium approach. The taxes examined in the study were export tax, import duty, excise tax, sales tax, education tax, drainage charges levied by the federal government, and land tax levied by the state government. INCIDENCE ASSUMPTIONS It was assumed that land based taxes 4 were borne by landowners, the statutory tax payers. In practice taxes on agricultural land are rarely shifted to consumers. In most developing countries shifting of land tax to tenants is not likely as the amount of rent paid by tenants and tenants' share of the cost of production are inflexible, determined mainly by noneconomic factors such as custom and tradition. Leases are generally long term and the relationship between landlord and tenant is more personal than businesslike (Wald, 959, p. 9). The legal incidence of an export tax is on the exporter from whom the government collects the tax. Under competitive conditions shifting of export tax forward or backward depends on the ratio of the elasticity of supply to the elasticity of demand as given by the Dalton (954, p. 5) formula: dp E s t (E s + E d ) where dp = increase in price t = amount of the tax E s = elasticity of supply Ed = elasticity of demand The demand for natural rubber which is determined by a host of technoeconomic factors, is believed to be price elastic. Tan (967, p. 96), for example, suggests that the price elasticity of demand for Malaysian natural rubber is 5. In the short run, the supply of natural rubber is generally inelastic because of its fixed productive capacity. Rubber takes six to seven years from the time of planting to the time of first tapping. An additional f\\t to ten years are required before MOHAMMED ARIFF BIN HUSSEIN peak yield is obtained. Although changes in the supply can be effected through changes in tapping frequency, size and number of tapping cuts, their effects are marginal 5 (Allen, 972, p. 7). Estimates of price elasticity for smallholders' productions have ranged from.8 to.37 (Behrman, 97; Wharton, 963; Chow, 976). Since most of domestic production is exported, the supply elasticity for export would also probably fall within that range. The interchangeability of coconut oil with other vegetable oils is extensive particularly in the manufacture of margarine and shortening. Different kinds of fat and oil are to a large extent substitutes since their characteristics can be altered by processing. Competition is also found between the natural products and their synthetic counterparts, especially between soap made from natural fats and synthetic detergents based largely on petroleum derivatives. These conditions would tend to make the demand for coconut oil very elastic. The supply of coconut by smallholders is also believed to be price inelastic. Like rubber the period between planting and the first harvest is about six years. Besides, coconut producers have less flexibility in terms of changing their rate of output under different price conditions. Unlike rubber which can be tapped daily, coconuts are harvested only once in every two months. Substitution effects in the short run are therefore relatively insignificant. Malaysia is only one of several producers of canned pineapple for the world market. Competition occurs not only among producing countries and also between pineapple and other tropical crops. One would expect, therefore, that the demand facing Malaysian products to be price elastic. Pineapple is a semipermanent crop. Tha economic life of pineapple in Malaysia is about ten years and the period between planting and the first harvest is 8 months. Pineapple growers, therefore, can easily adjust: their output in response to price changes. In addition, a substantial proportion of the crop is planted as a catch crop between rows of immature rubber trees where substitution with other catch crops is easily made. However, in the face of a very elastic world demand the tax burden is believed to be shifted back to producers. 4 L ot r? land d f\ d lots educati owned. n taxes with drainage changes are land based taxes as their rates are based on the size 5 Although ethrel stimulation has boosted elasticity somewhat, its application bv traditional rubber smallholders is still rather limited. 6

50 Under competitive conditions, import taxes will be shared between the seller and the buyer according to the respective elasticities of demand and supply 6 : Backward shifting of import tax to foreign producers, however, is highly unlikely since the quantity imported in the country is a small part of the total world import of any one particular commodity. From the point of view of the Malaysian consumer therefore, the supply of an imported commodity can be assumed to be perfectly elastic. Generally import taxes are regarded by importers as an addition to costs of supplying the imported goods to consumers. Therefore the full amount of the tax is passed on to consumers. Shifting of the tax burden forward to consumers is further enhanced by the practice of percentage markups on the part of importers. Eleven to fourteen percent markups are normally added to the import price plus duty (Edwards, 97, p. 62). As in the case of import duties, excise and sales taxes represent an addition to production costs and tend to be reflected in higher prices. The burden of an excise tax is therefore generally assumed to be fully shifted to consumers. The implicit assumption seems to be that of a perfectly elastic supply rather than a perfectly elastic demand. Most studies also assume that the sales tax is fully shifted to consumers. The assumption is that the supply of money capital is perfectly elastic and if market is not perfectly competitive, the firm will regard the tax as an addition to the costs of capital goods. The tax will then be reflected in higher prices of the consumption goods produced. Tax pyramidying which occurs when sales tax is levied at nonretail level further reinforces shifting of the tax to consumers. 7 INCOME ESTIMATES Secondary data sources were used to develop estimates of alternative income measures and total taxes paid by the smallholders. Income estimates for rubber smallholders were computed from data on average holding size, yield, price and costs of production. Esti TAX BURDEN ON JOHORE SMALLHOLDERS mates of average size were obtained from the 96 Census of Agriculture 8. The proportion of immature rubber acreage was based on all replanted and new planted acreage during a period of seven years ( ), while average yield was based on production figures of smallholders participating in group processing centres (GPCs). It was also assumed that the smallholders produced RSS 3 and 4 grade rubber. The price that they received was computed by deducting a 9 per cent marketing margin from RSS F.O.B. price less export duty and cesses (Lim, 968). Scrap rubber was assumed to constitute 5 percent by weight of the sheet rubber production. Cost of production estimates adjusted to take into account changes in price level was based on rubber smallholders who were members of GPCs (Bevan, 962; Barlow and Chan, 969). It was assumed that labour was provided only by family members and that economies of scale were absent within the range of holding size considered. The 96 census of agriculture and three specific surveys (Wilsons, 958; FAO, UN, 968; FAMA, 973) on coconut smallholders in West Johore were the main sources of data used to develop income estimates for coconut smallholders. As in rubber, estimates of the average size of coconut smallholdings in the state were based on the Census data. Comparable estimates from the four studies tend to corroborate the assertion that average size of holding has remained fairly constant. The FOA estimates of the palm density was used as it reflects the conventional belief that the palm density on smallholdings tend to be higher than what is recommended because of the tendency for farmers to let the nutfalls grow as a source of additional income. The proportion of bearing palms and equivalent estimates of palm bearing status were based on the FAMA survey. The FAMA survey also provided the most recent estimates of yield per acre. 6 Some researchers argue that the burden of import taxes can be ignored if they are levied primarily to discourage imports rather than to increase government revenue (Pechman and Okner, 974 p. 7). In Malaysia import duties are imposed mostly for revenue purposes (Edward, 97, p. 74). 7 Malaysia introduced an ad valorem single stage sales tax of 5 per cent on import and locally manufactured goods early in 972. * Changes in holding size are believed to be insignificant because of the low rate of growth in total acreage and the absorption of additional acreage by new entrants to agricultural labor force (Tan, 975, pp. 89). That the census estimate had been biased downward (Greenwood Word, 964) is believed to be of little significance since the study is only concerned with holdings less than 5 acres. 7

51 As the three surveys were confined only to the west coast districts of the state, estimates of palm density, bearing status and yield for the other districts were based on the census data 9. The FAMA survey shows there was little variation in the prices of husked nuts and copra between districts. The average monthly exfarm prices for husked nuts and copra for the twelve months of 975 as reported by FAMA were used to compute the alternative measures of income. The majority of coconut products were sold in the form of copra and husked nuts. Costs of producing coconut which depend on the type of coconut product sold were based on Selvadurai's (968, Table 73) estimates for the district of Batu Pahat and Pontian. The census of agriculture did not analyse pineapple as a separate crop. Estimates of pineapple production and income were drawn from surveys conducted by the Ministry of Agriculture (Selvadurai and Jegathesan, 968; Selvadurai etal, 975). Only pineapple of the canning variety grown in the district of Pontian, Batu Pahat, Kluang and Muar were considered. The price used to estimate gross returns was obtained through adjustment made on the fixed price offered to producers. The adjustment factor was the spread between the fixed price and the average price received by farmers in 974. Selvadurai's estimates of production costs in 974 were adjusted to take into account changes in the price level. ESTIMATES OF TAXES PAID The amount of land tax paid by smallholders depends on the size of alienated lots, whether the lots are being replanted, and whether they are in Malay reserve areas (Hussein, 977, Appendix 2). To obtain the estimates of alienated acerage for smallholders, the total acreage actually planted in estates and land schemes was subtracted from MOHAMMED ARJFF BIN HUSSEIN the total acreage alienated for each of the three crops. The alienated acreage estimated was then compared with the total acreage actually planted with the crop. In the case of rubber, the planted acreage was found to exceed the alienated acreage. It was assumed that the excess acreage was planted on land not alienated for rubber. Crops other than rubber were then assigned to the excess acreage based on the proportion of total acreage alienated for crops other than rubber. In the case of coconut, the planted acreage in smallholdings was less than the total acreage alienated for it. It was assumed, therefore, that coconut in smallholdings are planted on lots alienated for coconut. The registration record of the Malaysian Pineapple Industry Board (MPIB) provides detailed information on types of lots where pineapple of the canning variety is grown. This information was used to compute the estimates of taxes paid by pineapple smallholders. The size distribution of replanted rubber lots in the state from 952 to 974 was used as a proxy for the size distribution of lots alienated for rubber. The Coconut Smallholders Development Scheme keeps a record on the size of individual lots whose owners have applied for replanting or rehabilitation subsidy. The size distribution of these lots was used to categorise lots alienated for coconut. The size distribution of lots cultivated with pineapple as provided by the MPIB was used for the same purpose. Land tax rates for lots planted with rubber or coconut are lower for the first six years following approval for replanting or following alienation. The sixyear period corresponds approximately to the time taken for rubber and coconut to mature. Pineapple smallholders are eligible for the reduced tax rate during the first two years following replanting. The land tax for all countryland, ten acres or less, situated in Malay reservation where the owner or owners are Malays, is half the specified rate. The education tax at one ringgit per acre is levied on owners of all lots greater than three acres 2. Coconut acreage in the nonwest coast districts is only 3 per cent of the total smallholding acreage in the state. Except for the district of Mersing, replanting and rehabilitation programmes are concentrated exclusively on the west coast districts. The replanting and rehabilitation programmes in Mersing started in 973. It is conceivable that coconut is also being cultivated on land alienated for other crops. For example, Wilson (958, Table 7) shows that about 2 per cent of the coconut area in West Johore was cultivated on land not alienated for the crop. The assumption that all the coconut in the state is cultivated on land specifically alienated for the crop is incorrect to the extent that the above is true. The proportion of newly alienated lots in the traditional smallholdings is small because the state of Johore closed the land register for private applications in 96 and concentrated on group alienation in land development schemes (Guyot, 97, p. 384). 2 Lots three acres or less alienated for oil palm, however, are not exempted. 8

52 Drainage charges at $6 an acre are levied on coconut areas where drainage schemes are provided by the government. In Johore these schemes are located in the West Coast districts. All types of rubber exported from Malaysia are subject to export duty, surcharge, replanting cess, and research cess. The export duty and the surcharge vary directly with the prevailing market price of RSS (Hussein, 977, Appendices 3 and 4). The replanting and research cesses are levied at a flat rate of 4J cents per lb. and cent per lb. respectively regardless of the price level. The amount of export duty, surcharge, and cesses paid by rubber smallholders were determined from estimates of total production less local consumption and the prevailing price of RSS. The main coconut products exported are fresh nuts, copra and coconut oil. To encourage domestic processing of coconuts into coconut oil, export duty is levied on fresh coconuts and copra at a rate of per cent ad varolem. The total tax revenue collected was computed from the total value of fresh nuts and copra exported. On the basis of planted acreage and yield, the tax revenue was allocated between estate and smallholding sector, among coconut producing states and between smallholders of less than 5 acres and those of 5 acres or more. An export cess is levied on canned pineapple exported from Malaysia. The rate, which is fixed per unit of export, varies according to importing countries (Hussein, 977, Appendix 7). The total amount of the cess revenue collected was computed from the total number of standard cases exported to the respective countries. The total amount was then allocated evenly between producers and canners 3, between the estate and the smallholding sector based on production shares. The amount accrued to the smallholding sector was further divided between smallholders of less than 5 acres and those of 5 acres or more. Except for formic acid, the other inputs used in rubber production are generally not subject to import, excise and sales taxes. An import tax of 25 per cent ad valorem is levied on formic acid. In the case of coconut, the cost items generally subject to these taxes are the equipment, implements and tools. The amount of these taxes paid by smallholdings was determined from the tax rates and from estimates of the average annual depreciation on the items. TAX BURDEN ON JOHORE SMALLHOLDERS Among the material inputs used in pineapple production, only weedicides and insecticides are subject to import duties. The equipment and machinery are generally subject to at least one of the three taxes. As in rubber and coconut, the amount of the three taxes paid was determined from estimates of the total value consumed and from the appropriate tax rates. RESULTS AND CONCLUSIONS Table gives the amount of taxes paid by the smallholders. The most important category of taxes borne by them is shown to be export based. Export duty, surcharge, and export cesses account for 86 per cent of all taxes paid. Land tax which is an important source of revenue to the state government accounts for only eight per cent of the taxes. The export based taxes are also the most important taxes paid by rubber and pineapple smallholders. Respectively they account for 9 per cent and 48 per cent of all the taxes paid. The most important category of taxes paid by coconut smallholders are landbased, notably the drainage charges which account for 52 per cent of the total taxes paid. Table 2 gives the estimates of tax burden. For the alternative income measures used, rubber smallholders pay relatively, higher taxes than either coconut or pineapple smallholders. Depending on the income measures used, the average rubber smallholder pays onefourth to onethird of his income in taxes. If the quid pro quo taxes are not considered, the tax burden on the smallholders will be reduced to about onehalf the amount when all taxes are considered. However, the burden on rubber smallholders is still the highest. The possibility of increasing real income of smallholders by eliminating the taxes paid is limited. Export cesses, education tax and drainage charges are levied for specific purposes. Eliminating or reducing them would adversely affect the functioning of the respective agencies, programmes or projects. Smallholders as purchasers of production inputs are already being exempted from paying import, excise and sales taxes for most of the items they consume. For many inputs where such taxes are levied they are also being consumed by other sectors of the economy. 3 This assumption is based on the fact that the price fixing is done through a bargaining process where the interests of producers and canners are equally represented. 9

53 TABLE Estimated Taxes Paid by Rubber, Coconut and Pineapple Smallholders of Less Than 5 Acres; Johore, 975. Type of Smalholding Rubber Land Tax (8) 2,78,39 Education Tax (S) 384,98 Drainage Charges ($) _ Export Duty and Surcharge (W,556,378 Export Cesses 6,42,6 Import Excise and Sales Taxes (S) 744,44 Amount (S) 3,65,63 Total Percent 95.7 Coconut 432,66 88,49 62,496 3,93 25,2,98, Pineapple Total 33,7 9,69 :V 88,775 53,369 85,464.6 X Amount Percent 2,544, , ,496.9,588, ,49, , ,549,8. m D.V t Smallholder Total Area in Smallholdings, Total Area of Smallholdings Less than 5 Acres (acres) Average Size of Holdings (acres) TABLE 2 Average Size and Number of Holdings, Amount of Taxes, Income and by Type of Smallholding; Johore, 975. Total Number of Holdings Amount of Taxes Paid per Holding Income Net Gross A, (S) Net B' Tax Burden Gross Income Tax Bturden Index Net Income A' Net Income B' ARIFF BIN HIJSSEIN Rubber 59, , , Coconut 4, , Pineapple,37 6., , A* Net income after subtracting all costs except family labor from gross income. B* Net income after subtracting variable costs from gross income.

54 TAX BURDEN ON JOHORE SMALLHOLDERS Possible changes that could be made would include exempting smallholders of certain size groups from land tax and compensating the state government for the revenue loss by imposing progressively higher tax rates on larger holdings. For the export tax and surcharge, a system of rebate could be designed to benefit smallholders of certain size groups. Results of the study show that real income of smallholders especially that of rubber smallholders, is reduced substantially because of taxes. However, the scope for increasing income through changes in the tax system is rather limited. Programmes that increase farm productivity and size of holding should complement such changes. ACKNOWLEDGEMENT The author wishes to express his gratitute to Prof. J. Dean Jansma for his encouragement and guidance throughout the course of this study. REFERENCES ALLEN, P.W., (972): Natural Rubber and Synthetics. New York. John Wiley and Sons. BARLOW, C. and CHAN, C.K., (969): Toward an Optimum Size of Rubber Holding. J. Rubb. Res. Inst. Malaysia. 2, (5) BEHRMAN, JR., (97): Econometric Model Simulations of the World Rubber Market, Essays in Industrial Econometrics Vol. Ill, L.R. Klein (ed), Wharton School of Finance and Commerce, University of Pennsylvania. BEVAN, J.W.L., (962): A Study of Yields, Labor Inputs, and Incomes of Rubber Smallholdings in the Coastal Area of Selangor. Faculty of Agriculture, University of Malaya, Kuala Lumpur. CHOW, C.S., (976): Some Aspects of Price Elasticities of Rubber Production in Malaysia. Proc Int. Rubb. Conf. Kuala Lumpur, 975. DALTON, H., (954): Principles of Public Finance. London: Routledge and Kegan Paul. EDWARDS, C.T., (97): Public Finance in Malaya and Singapore. Canberra: Australian National University Press. FEDERAL AGRICULTURAL MARKETING AUTHORITY (FAMA), (973): Coconut Production and Marketing Survey of West Johore. GREENWOOD, J.M.F., (964): Rubber Smallholdings in the Federation of Malaya. J. trop. Geogr. 8, 8. GUYOT, D., (97): "The Politics of Land: Comparative Development in two states of Malaysia/* Pacific Affairs. An International Review of Asia and the Pacific 55(3). HUSSEIN, M.A., (977): Tax Burden on Rubber, Coconut and Pineapple Smallholders in Johore, Malaysia. Unpublished Ph.D. thesis. Pennsylvania State University, U.S.A. LIM, S.C, (968): A Study of the Marketing of Smallholders Rubber at the First Level in Selangor. Kuala Lumpur: Economic and Planning Division of the Rubber Research Institute of Malaya, Report No. 2. MCLURE, CE. Jr., (972): The incidence of Taxation in West Malaysia, Malayan Economic Review, 7. PECHMAN, J.A. and OKNER, B.A., (974): Who Bears the Tax Burden. Washington, D.C: The Brookings Institution. SELVADURAI, S, (968): Preliminary Report on the Survey of Coconut Smallholdings in West Malaysia, Ministry of Agriculture and Cooperative* Kuala Lumpur. ABDULLAH BIN MOHD. YUNOS, YAP KIM LIAN, and LEE KIM SEONG (975): Socioeconomic Survey of Pineapple Smallgrowers in Johore. Ministry of Agriculture and Rural Development, Kuala Lumpur. SELVADURAI, S. and JEGATHESAN, S., (968): An Economic Survey of Pineapple Smallholdings in Pontian, Johore. Ministry of Agriculture and Cooperative, Kuala Lumpur. SNODGRASS, D.R, (975): "The Fiscal System as an Income Redistributor in West Malaysia". Readings in Malaysian Economic Development. David Lim (ed). Kuala Lumpur, Oxford University Press, TAN, A.A.H. (967): The Incidence of Export Taxes on Small producers. Malayan Economic Review. 2. TAN, B.T. (975): "Agricultural Subsidy and Price Support Policy.'* Agenda for the Nation, Second Malaysian Economic Convention, Kuala Lumpur: Malaysian Economic Association. UNITED NATIONS FOOD AND AGRICUL TURAL ORGANISATION (968): Rome: Economic Survey of the Coconut Growing Industry: Report to the Government of Malaysia. WALD, H.P., (959): Taxation Agricultural Land in Underdeveloped Economics: Cambridge: Havard University Press. WILSON, T.B., (958): The West Johore Coconut Production Survey. Ministry of Agriculture, Kuala Lumpur. (Received 8 August 978)

55 Pertanika (2), 29 (978) Floristic Components of the Ground Flora of a Tropical Lowland Rain Forest at Gunung Mulu National Park, Sarawak RUTH KIEW Department of Biology, Faculty of Science and Environmental Studies, Universiti Pertanian Malaysia Key words: Floristic Components, Ground Layer, Lowland rainforest, Microhabitat, Distribution of tropical Herbaceous plants, Variegated foliage. RINGKASAN Komponen tumbuhtumbuhan lapisan tanah yang berkiatan dengan mikrohabitat di Taman Negara Gunung Mulu Sarawak telah dihuraikan. Mikrohabitat pada sistem sungai merangkumi kawasan air cetek, tebing berbatubatu, tebing sungai luas, dan mikrohabitat dihutan teduh iaitu kawasan selalu lembah, lereng berbatu tanpa sarap, lereng curam bersarap, hutan lanar rata tersalir, permukaan kayu balak, akar dan batu dapat disiasat. Punca sebab yang mungkin menyebabkan penyibaran yang tak sama rata spesiesspesies dilapisan tanah hutan lanar rata tersalir telah dibincangkan. Senari spesies hutan lanar inijuga telah dibandingkan dengan senari spesies yang telah dilapurkan dalam penerbitan dari Lembah Danum Sabah, Hutan Simpan Pasoh Malaysia Barat dan lain hutan pamah di Sarawak, seperti kerangas dan hutan paya gambut. Kehadiran daun berwarna pelangi dan beranika warna telah dicatitkan dengan ringkas. SUMMARY The floristic components of the ground layer at the Gunung Mulu National Park, Sarawak, are described. These components are associated with microhabitats. The microhabitats of the riverine system include the shallow stream, rocky banks and banks of larger rivers, and those of the shaded forest include permanently wet areas, rocky litterfree slopes, steep litter covered slopes, surfaces of logs, roots or rocks and the flat welldrained alluvial forest. Possible causes of the uneven distribution of the ground layer species of the flat welldrained alluvial forest are discussed. The species composition of this alluvial forest is compared with other published reports from the Danum Valley, Sabah and Pasoh Forest Reserve Malaya, and with some other lowland forest types such as ke ran gas and peat swamp forest in Sarawak. The phenomenon of iridescence and variegation of leaves is briefly noted. INTRODUCTION The ground flora of the tropical rain forest (i.e. plants below 2m tall) lives in one of the most uniform environments, characterised by extremely low light intensity (about % of full sunlight), high humidity (9,,) and constant temperatures and where the diurnal difference is greater than the annual. In spite of these constant environmental factors and the fact that many species have a wide geographic range, their distribution within the rain forest is by no means uniform. On the one hand, microhabitats can be distinguished that possess a distinctive floristic composition: on the other hand the distribution of plants of the flat alluvial forest shows no obvious pattern of distribution and large areas of the forest floor are devoid of herbaceous cover. Information on the natural history and distribution of species of the ground flora of Malesian 2 forests is limited to the local floras, except for Burtt's review (977) on growth patterns and reproductive methods of the herbaceous flora. Although many ecological surveys of forest types have been carried out in Malaysia, these concentrate on the larger trees and information on the ground flora is often lacking. The survey described here is based on five week's fieldwork in the lowland alluvial forest of the Gunung Mulu National Park, Sarawak (Fig. ). General descriptions of the ground flora of tropical rain forest (Richards, 952, Burtt, 977) mention the predominance of certain families: Araceae, Begoniaceae, Cyperaceae, Gesneriaceae and Melastomaceae (at Mulu, Acanthaceae and Rubiaceae are also common) and comment that the number of families and species involved is very much less than those of the tree layers, probably due to the less favourable conditions for plant growth imposed by the deep shade;

56 BRUNEL Fig.. Map of Sarawak to show the location of the Gunung Mulu National Park (M). but that several genera are obviously successfully adapted to these conditions and possess many species e.g. Begonia (Begoniaceae)and Cyrtandra (Gesneriaceae). While these families are herbaceous, there is also an important component of the ground flora which is woody, i.e. the diminutive palms (in particular the genera Pinanga, Iguanura and Licuala) Euthemis (Ochnaceae) and Driessenia (Melastomaceae). Apart from the angiosperm flora, ferns and Selaginella are important, though there is a noticeable absence of ground mosses as compared (for example) with kerangas [ forest where they form a conspicuous feature of the ground flora. Floristic components of the ground flora Within the lowland forest at Mulu several microhabitats can be distinguished which possess a characteristic assemblage of species. Broadly these can be grouped into those in which the canopy is open (Table ), for example, along streams; in contrast with those in which the canopy is closed (Table 2). Within the undisturbed forest, the canopy over streams is sometimes not complete and direct sunlight can reach a restricted area of the forest floor. Two such streams were observed in the study area (Table A) where the characteristic species include Alocasia aff. macrorrhiza (L.) Schott. which is aquatic and also appears lightdemanding; and Curculigo latifolia Dryand., which although relatively lightdemanding, requires better drained soils and thus is common on streambanks. The steep, almost vertical, rocky banks of torrential streams carry a characteristic flora (Table IB). A 5m long portion of the bank on the Sungai Tabaun (Fig. 2) was sampled intensively and the general results for RUTH KIEW species composition and relative abundance were compared and confirmed by observation along the Melinau Gorge. The lower part of the rocky bank was subjected to intermittent flooding after rain, and here Piptospatha elongata N.E. Brown was abundant (and confined to this habitat). Also present was a species of Sonerila and small plants of Myrmeconauclea strigosa (Korth.) Merr., which in this habitat did not appear to grow to adult size or to flower. Above the flood level, M. strigosa was the commonest plant, with its characteristically horizontal branches reaching over the river. In addition there were eight other species of flowering plants (Table B), seven species of ferns and one species of Selaginella. Above the rocky bank at Sungai Tabaun, towered the damp shaded vertical rock face of the gorge which was completely covered by plants of Hexatheca fulva C.B.C., many with snowy white flowers. Epithema involucratum (Roxb.) Burtt occurred on rocks in the river. Another riverine association was found along the banks of the larger rivers especially on banks which were wide and where small boulders had accumulated. These observations were made below the Melinau Gorge (Fig. 2). Here the shrub layer was very dense, frequently covered by a variety of flowering woody creepers (Table C) and a few r herbaceous climbers, such as species of Aeschynanthus (Gesneriaceae) and Hoseanthus lobbii (Verbenaceae). Only in open areas, e.g. where there was a clearing, were herbs able to flourish. Some of these species are lightdemanding and become relatively large: Musa campestris Becc, Donax canniformis (Forst.) K. Schum. and Costus speciosus (Koenig) J.B. Smith, for example, attain a height of 2m. However, the species diversity of the ground flora of this habitat is poor. Within the lowland forest itself, where the canopy is complete, several distinct microhabitats can be recognised by their characteristic assemblage of species (Table 2). These include a) permanently wet areas, b) rocky litterfree slopes, c) epiphytes on logs, roots or boulders, d) steep littercovered slopes and e) the flat alluvial forest. The wet areas probably never dry out, because at Mulu the rainfall is distributed evenly throughout the year (J. Procter, pers. comm.). They also are likely to be subjected to frequent water Jogging; about.5m of standing water was observed after a night of heavy rain in one such area. These areas are conspicuous for the dense cover of the ground layer, often with a single species dominating, such as Alpinia glabra Ridl. kerangas forest is defined by Brunig (975) as "Tropical lowland forests on infertile soil derived from basepoor parent material. 3

57 FLORISTIC COMPONENTS OF GROUND FLORA AT GUNUNG MULU, SARAWAK A. Shallow streams Acanthaceae: Staurogyne setigera Nees. O. Kunzte Araceae: Alocasia aff. macrorrhiza (L.) Schott. Begoniaceae: Begonia sp i Begonia sp. ii Campanulaceae: Pentaphragma acuminaia Airy Shaw Hypoxidaceae: Ciirculigo latifolia Dry and Urticaceae: Elatostema sesquifolittm (Bl.) Hassk. Ii, Rocky banks of torrential streams Araceae: Homalomena sagittifolia Jungh. Schott Piptospatha elongaia N. E. Brown Gesneriaceae: Epithema involucratum (Roxb.) Burtt Hexatheca fulva C.B.C. Marantaceae: Stachyphrynium sp. Melastomaceae: Sonerila sp. i Ochnaceae: Neckia serrata Korth. Rubiaceae: Argostemma sp. Myrmeconauclea strigosa (Korth.) Merr. Urticaceae: Elatostema sesquifolittm (Bl.) Hassk. C. Banks of wide rivers Shrubs (including small or young trees): Capparidaceae: Cratera nurvala Ham. Dilleniaceae: Dillenia suffruticosa (Griff.) Martelli Leeaceae: Leea aculeata Bl. TABLE Ground layer species associated with riverine systems. (* These families are under revision at Kew Gardens, London). or Hanguana malayana Merr. which are monocotyledons and spread by means of suckers or rhizomes. Iguanura melinauensis Kiew is frequent in this habitat and its seedlings must be able to withstand submersion. Polyalthia flagellaris (Becc.) AiryShaw, a small annonaceous tree with deep red flowers and fruits on short knobbly twigs above ground level, is conspicuous in this habitat. 4 Melastomaceae: Medinilla crassifolia Bl. (epiphyte) Piperaceae: Piper caninum Bl. Polygalaceae: Polygala venenosa Juss. Sonneratiaceae: Duabanga moluccana Bl. Rubiaceae: Myrmeconauclea strigosa (Korth.) Merr. Psychotria crassifolia Miq. Climbers: Convolvulaceae: Merremia peltata (L.) Merr. Cucurbitaceae: Trichosanthes trifoliolata V. Merell Gesneriaceae: Aeschynanthus parvifolius R. Br. A. tricolor Hook. f. Oleaceae: Jasminum crassifolium Bl. Rubiaceae: Uncaria sp. Smilacaceae: Smilax odoratissima Bl. Verbenaceae: Hoseanthus lobbii (Cl.) Ridl. Vitex pubescens Vahl. Vitidaceae: Vitis cinnamomea Wall. Herbs: Araceae: AmorphophaUus sp. Gramineae: Centotheca lappacea (L) Desv. Marantaceae: Donax canniformis (Forst.) K. Schum. Musaceae: Musa campestris Becc. Palmae: Pinanga riparia Becc. Pandanaceae: Pandanus sp. Rubiaceae: Acranthera frutescens Val. ex Winkler Zingiberaceae: Costus speciosits (Koenig) J. E. Smith Rocky slopes, although small in area, support a specific ground flora (Table 2B) including species of Selaginella and Hymenophyllaceae. These slopes are steep, with a crumbling rock surface without leaf litter. The species composition appears uniform from the two areas observed (which were 8km apart) (Fig. 2), and several species such as Driessenia axanthera Korth,. Homalomena humilis var ovalifolia Hotta, and Phyllagathis elliptica Stapf appear to be exclusive to such areas. On rocky slopes in narrow gulleys, damper conditions prevailed and Begonia species were more conspicuous. Neckia serrata Korth., by contrast, is a common and very widespread species which is also found on rocky banks of streams, suggesting that it is tolerant of lighter conditions, but is apparently intolerant of an accumulating litter layer as it is also found on the forest floor clear of litter. Neckia serrata is atypical in this respect as no

58 RUTH KIEW \ v, x * r_; Fig. 2. Map of the Gunung Mulu National Park, Sarawak. (M.G. Melinau Gorge; # camps; R sites of rocky slopes and ridges). other species of this assemblage also grows by streams. The epiphytes of the ground layer of the lowland forest occupy two habitats: fallen logs, buttresses and raised tree roots or boulders, which at Mulu are generally limestone and which are frequently completely covered by herbs. Of these species (Table 2C) only Elatostema acuminata (Poir) Brongn. appears to be widespread and is found both on raised tree roots 5 and boulders. Cyrtandra oblongifolia (Bl.) C.B.C., a species of Sonerila and Lobelia zeylanica L. appear as small, solitary plants on fallen logs. The remaining species which were found on limestone rocks were larger plants and spread over the rock surface by rooting at the nodes. Limestone boulders are often completely covered by epiphytes probably because the depressions on their surface retain water and perhaps allow litter accumulation.

59 FLORISTIC COMPONENTS OF GROUND FLORA AT GUNUNG MULU. SARAWAK TABLE 2 Ground layer species associated with shaded forest habitats. A. Permanently wet areas Araceae: Homalomena insignis N. E. Br. Flagellariaceae: Hanguana malayana Merr. Palmae: Jguanura melinauensis Kiew Zingiberaceae: Alpinia glabra Ridl. Achasma megalocheilos Griff. Geanthusfimbriobracteatus (K. Schum.) Burtt & Smith B. Rocky, litterfree slopes Araceae: Homalomena humilis Hook. f. var. ovalifolia Hotta # Begoniaceae: Begonia sp. iii Begonia sp. iv Begonia sp. v Begonia sp. vi Melastomaceae: Driessenia axantha Korth. Phyllagathis elliptica Stapf Ochnaceae: Neckia serrata Korth. Rubiaceae: Argostemma borraginenm Bl. Triuridaceae: Sciaphila flexuosa Giessen C. Epiphytes on logs, roots, and boulders Acanthaceae: Hallieracrantha salicifolia Stapf *Begoniaceae: Begonia sp. vii Burmanniaceae: Burmannia championii Thw. Campanulaceae: Lobelia zeylanica L. Gesneriaceae: Cyrtandra incrustata (Bl.) Burtt C. oblongifolia (Bl.) C.B.C. Loxocarpus sp. Melastomaceae: Sonerila sp. ii Rubiaceae: Argostemma havilandii Ridl. OphiorrhizafibriUosa Ridl. Urticaceae: Dendrocnide stimulans (L.f.) Chew (shrub) Elatostema acuminata (Poir) Brongn. D. Steep littercovered slopes # Shrubs and small trees: Annonaceae: Goniothalamus malayanus Hook. f. Araceae: Alocasia denudata Engl. Gnetaceae: Gnetum gnemon var. tenerum Markgraf G. macrostachyum Hook. f. (Liana) Melastomaceae: Phaulanthus acuminatissimus Ridl. Myrsinaceae: Ardisia javanica DC Palmae: Eugeissona utilis Becc. Rubiaceae: Cephalis stipulaceae Bl. Herbs: These families are under revision at Kew Gardens, London Asclepiadaceae: Hoya parasitica Wall, (climber) Gesneriaceae: Cyrtandra bracheia Burtt C. pendulifera Kraenzl Loxocarpus verbeniflos (CB.C.) Burtt Marantaceae: Stachyphrynium sp. Melastomaceae: Sonerila pulchella Stapf Sonerila sp. iii Ochnaceae: Euthemis leucocarpa Jack Neckia serrata Korth. Palmae: Areca tenella Becc. Licuala sp. Pinanga tomentella Becc. P. disticha (Roxb.) Wendl. Pinanga sp.ii Pinanga sp. iii Pinanga sp. iv (variegated) Rubiaceae: Argostemma psychotrioides Ridl. Zingiberaceae: Alpinia sp. i Alpinia sp. ii Boesenbergia sp. Plagiostachys sp. H. Flat, welldrained alluvial forest Acanthaceae: Cosmianthemum magnifolium Bremekamp Staurogyne setigera Nees. O. Kuntze Campanulaceae: Pentaphragma cyrtandriforme Airy Shaw r Commelinaceae: Forrestia marginata Hassk. Cyperaceae: Mapania cuspidata (Miq.) Uttten Gesneriaceae: Cyrtandra radiciflora CB.C. Marantaceae: Stachyphrynium sp. Phrynium capitatum Willd. Orchidaceae: Calanthe triplicate (Willem.) Ames Malaxis aft. nemoralis (Ridl.) Holttum Plocoglottis art. javanica Bl. Palmae: Iguamira melinauensis Kiew Piperaceae: Piper porphyrophyllum N.E.Br. Polygalaceae: Epirixanthes cylindrica Bl. E. elongata Bl. Rubiaceae: Acranthera involucrata Val. Argostemma psychotrioides Ridl. Lasianthus stipularis Bl. (small shrub) Myrioneuron cyaneum Hall. f. Ophiorrhiza communis Ridl. Streblosa bracteata Ridl. Taccaceae: Tacca sp. Triuridaceae: Sciaphila flexuosa Giessen Urticaceae: Elatostema acuminata (Poir) Brongn. E. variolaminosum Schroeter Zingiberaceae: Boesenbergia pulchella Ridl. Geostachys sp. Globba atrosanguinea T & B 6

60 RUTH KIEW The assemblage of plants found on litter covered steep slopes and ridges (Table 2D) has few species in common either with rocky slopes (where litter does not accumulate) or with the low undulating alluvial forest floor. This is particularly true of the gingers which appear to have quite strict requirements for water. Exceptions are Argostemtna psychotrioides Ridl. and Stachyphrynium (which also grow in the flat alluvial forest) and Neckia serrata Korth. (which grows on shaded rocky slopes and exposed rocky stream banks). Richards (936) noted that there was a difference in the abundance of herbs on ridgetops (four times as many) than on slopes, which he suggested could be due to the more open canopy of ridge tops increasing the light intensity at ground level. From observation at Mulu, undergrowth palms are more abundant on the ridges than on the slopes. However, some species such as Cyrtandra pendulifera Kraenzl and Sonerila pulchella Stapf, which appear as dense though small populations, appear specific to steep slopes. It is difficult to explain the differences between the two ridges at Mulu (Fig. 2). One supported a varied flora of about fourteen herbaceous species (Table 2D) while the other at Sungai Berar supported a few sterile plants of Stachyphrynium, Pandanus and gingers; though both supported a rich palm flora. Both ridges were apparently similar in slope, the only significant difference being that the latter ridge was shale. The flat welldrained alluvial forest covers a greater area than any of the other microhabitats described above and its flora is correspondingly diverse (Table 2E). Compared with the habitats described above, the plants are in general less frequent and give an impression of a large area of bare ground. A few of these species (Table 2E) such as Cyrtandra radicifolia C.B.CL. and Lasianthus stipularis Bl. form dense stands in particular areas. It is very difficult though to discern any difference between the areas where they occur and the areas from which they are absent. This is also true in the case of other species which show a patchy distribution. Some are represented by individuals, such as the ground orchids or species of Acranthera, while others are found as isolated populations, such as Ophiorrhiza communis Ridl., Argostemma psychotriodes Ridl. and Streblosa bracteata Ridl, Distribution of the ground flora of the alluvial forest The patchiness of the ground flora of the flat welldrained alluvial forest has been ascribed to three causes: the predominance of vegetative reproduction (Richards, 952, Walters, 955), the lack of pollination (van Steenis, 969) and the failure of seedling establishment (Burtt, 977). 7 Richards (952) considered species growing under the closed canopy as shadeloving and suggested that vegetative reproduction was more important in those species than reproduction by seed. In this survey there was no evidence for this claim as the great majority of the species could be found flowering or fruiting. The main exception was the Araceae. Many aroid species though frequently sterile, occur as solitary plants, which suggests that they are the product of seed dispersal. Burtt considers that vegetative reproduction is the cause of gregariousness and contrasted the vegetative spread in monocotyledons by suckers or rhizomes with that of the dicotyledons where clumps result because the tall aerial stems eventually become decumbent, root and produce aerial shoots. This is supported by the observations in this survey. These gregarious species (Argostemma, Streblosa and Elatostema) were as free flowering as the solitary species, such as Acranthera, Neckia and Sonerila. Insufficient pollination was suggested by van Steenis (969) as a limiting factor in the distribution of basally flowering trees, such as Apocynaceae and Bignoniaceae based on observations that they rarely set fruit. This is not true for the ground layer. Careful examination of plant populations in the field shows that most species produce fruits but they often tend to be inconspicuous and thus may be overlooked. One notable exception from Malaya is Lepidagathis longifolia Wight which Ridley (923) quotes as being abundant but "no one as yet has ever seen a fruit, and it would not fruit in Singapore Gardens". Henderson (959) mentions that none of the specimens of this species in the Singapore Herbarium had fruits. Many of the species observed at Mulu possessed both flowers and fruits on a single plant suggesting that flowering is more or less continuous. Failure of seedling establishment is difficult to assess, because seeds collected in the field and germinated in pots often show high viability (Kiew, 972, Burtt, 977), whereas in the field, seedlings are often rare. The most conspicuous seedlings in the ground layer are those of various species of palms, most of which belong to Lepidocaryoid rattan genera but arecoid species are well represented. The undergrowth palms are often solitary and rely on sexual production for survival. Perhaps the infrequency of seedlings of other species indicates that seedling establishment is the critical phase in their life history. In this case, vegetative reproduction can be viewed as a method of prolonging the life of the individual plant, thereby increasing the quantity of seed that a single plant can produce and so increasing its chances of replacement. The mode of dispersal

61 FLORISTIC COMPONENTS OF GROUND FLORA AT GUNUNG MULU, SARAWAK of the small dry seeds of these dicotyledonous herbs such as Gesneriaceae, Begoniaceae, Rubiaceae and Melastomaceae is not known, Burtt suggests that dispersal is effected by rainwash or by being carried on the feet of animals (wind dispersal is impossible, since wind movement in the undergrowth is negligible). However, in conditions outside the forest similar sized seeds belonging to rubiaceous weed genera such as Borreria are effectively dispersed as is evidenced by their abundance and rapid spread since their introduction, e.g. B. alata (Aubl.) DC. (B. latifolia) first noticed in Singapore in 95 (Ridley, 923) is now a widespread weed in Malaya. It is not known how its seeds are dispersed. In this survey, Iguanura melinauensis Kiew, was found to be a conspicuous example of a plant with a local distribution: it is common along the Sungai Melinau but is absent from the adjacent Sungai Berar. Many of the undergrowth species show widespread geographic distribution, which indicates that in the long term their mode of dispersal is effective. However, their patchy distribution within the undergrowth and the dearth of seedlings suggest that the seedling establishment phase is the critical one in most cases. The reason for this is not known. Perhaps predation of fruit might account for the lack of seedlings, rather than attack of the seedlings; most adult plants appear free of fungal and insect attack with the notable exception of Streblosa bracteata Ridl. which has most of its leaves perforated due to insect damage in the bud. Competition with tree roots for water in the soil surface might also be a limiting factor as the only closed herbaceous communities are found in areas which are permanently wet. There are a few species with an apparently scattered and widespread distribution, though they are nowhere common, e.g. Globba atrosanguinea T & B, Epirixanthes elongata Bl. and Acr anther a involucrata Yah. The distribution of a few species is correlated with soil conditions, as mentioned above. Neckia serrata requires a litterfree habitat, and Mapania cuspidata (Miq.) Uttien grows on more acid soils. The majority of species, however, have distributions which apparently have been determined more by chance. Comparison of ground flora at Mulu with other areas and forest types Comparison of the ground flora at Mulu with other areas is difficult because whereas the ground flora at Mulu has been well collected over the years, very few other areas are equally well known or published information is lacking. More significant is the absence of certain species 8 which are common in other areas. The Danuui Valley (Sabah) expedition botanical report on ground flora (Kiew, 977) based on B.C. Stone's collections, shows a close similarity to that as Mulu and is particularly useful for its descriptiont of the ridge, valley, riverine and gulley habitats: the species composition matches that of Mulu closely. For Malaya, no detailed studies of the herbaceous flora are reported. Soepadmo and Kira (977) in a brief summary of common families and genera of the ground flora at Pasoh F. R., Malaya, include Dracaena (correctly Pleomele) (Agavaceae), Phyllagathis (Melastomaceae) and Labisia (Myrsinaceae) which are common plants of the Malayan undergrowth. Species of Phyllagathis and Labisia were not collected from the Danum Valley, and neither is common at Mulu, whereas several species of Phyllagathis are abundant in the Malayan alluvial forest. At Mulu the one species of Phyllagathis collected was confined to rocky litterfree slopes. In contrast, species of Acranthera, which are widespread in Borneo, are not recorded from Malaya (Ridley, 923). However it is not unknown for different species of a genus to behave differently in separate localities, as in Iguanura (Kiew T, 976) where./. wallichiana in Malaya is common and widespread while in Sarawak no Iguanura species is widespread and many have extremely local distributions, c.f. /. melinauensis. Comparison of species diversity in this Mulu lowland forest ground flora with other forest types is difficult. This study is based on observation of about 5km 2, while the detailed studies of Brunig (974) on kerangas forest and Anderson (963) on peat swamp forest include data from all available sites in Sarawak. The 5 species of the ground flora collected at Mulu is an underestimation for the total ground flora of Sarawak as the exclusion of unidentified sterile plants underrepresents the Araceae, Orchidaceae and Zingiberaceae, and those species with a local distribution outside this study area were not included. Even so, the species diversity is very much higher than that of kerangas forest (69 species) and peat swamp forest (9 species). Compared with these two forest types, the alluvial forest is rich in species of Gesneriaceae, Begoniaceae (Begonia) and Urticaceae (Elatostema) but is poor in Nepenthaceae (Nepenthes), which is well represented in the other two forest types. The kerangas resembles the alluvial forest in its Melastomaceae, Rubiaceae, Palmae and Zingiberaceae which were well represented. In contrast there are only three species of Araceae, although the Cyperaceae with eleven species is better represented. The ground flora of the peat swamp forest can be expected to be low in species as much of the area is permanently submerged.

62 Several characteristic families of the ground layer of alluvial forest such as Acanthaceae, Melastomaceae, Orchidaceae and Rubiaceae are absent, none the less the Araceae is as well represented as in the alluvial forest though by different species. The lower species diversity of the kerangas and peat swamp forest can be ascribed to their environment offering less favourable conditions for plant growth. Kerangas forest is poor in nutrients and has a low ph and the swamp conditions of the peat swamp forest exclude those plants that require well drained soil. The incidence of iridescent and variegated foliage Most authors comment on this conspicuous feature of the ground flora noting that it is not seen in other layers of the forest. In fact very few species are involved and these are confined to even fewer families. Iridescence is common in some species of Selaginella, e.g. S. willdenowii, in which Lee and Lowry (975) showed that the iridescence was due to lensshaped epidermal cells which they suggested are advantageous in absorbing a wider spectrum of light intensities inthe deep shade. Iridescent foliage is also known in some ferns, some begonias and species of Mapania, Cyperaceae (Burtt, 977). Plants with variegated leaves (with white, yellow or reddish blotches, spots or lines) are more common than plants with iridescent foliage. At Mulu, genera with variegated leaves included Begonia, Sonerila, Alocasia, Plocoglottis, Malaxis, Pinanga, Piper and Pleomele (the latter collected by Stone, pers. comm.). Some of these species were constant in their variegation, such as Plocoglottis javanica Bl. with its white spots and Malaxis nemoralis (Ridl.) Holttum with its gingercoloured leaves; others such as Sonerila pulchella Stapf occurred as local populations and varied in the proportion of individuals with variegated leaves. Hibbert (87) illustrates cultivated examples, several of which belong to the Acanthaceae. Although herbaceous species of the Acanthaceae are common in the ground layer, relatively few species are variegated, as for example Gymnostachyium magisnervatum C.B.C. and Polytrema cupreum Ridl. from Malaya but no variegated examples of this family were collected at Mulu. Some of these species undoubtably have horticultural potential, in particular the begonias. Dransfield (974) has introduced several species of variegated pinangas (where the variegation takes the form of an attractive marbling of the leaves) into cultivation at the Bogor Botanic Gardens. Trial cultivation is necessary for the other species to determine their horticultural value. RUTH KIEW 9 ACKNOWLEDGEMENTS I wish to express my gratitude to the Royal Geographical Society for permission to participate in the Gunung Mulu (Sarawak) Expedition; to Universiti Pertanian Malaysia for the research grant which enabled me to participate; to Mr. Paul Chai, Forest Botanist, for providing facilities at the Forest Department, Kuching and to Dr. B.C. Stone for his helpful criticisms of this paper. REFERENCES ANDERSON, J.A.R. (963): The Flora of the Peat Swamp Forest of Sarawak and Brunei. Gdns* Bull Singapore 2: BRUNIG, E.F. (975): Ecological Studies in the Kerangas Forest of Sarawak and Brunei. Borneo Lit. Bureau, Govt. Printer, Kuching. BURTT, B.L. (977): Notes on RainForest Herbs. Gdns' Bull Singapore 29: 738. DRANSFIELD, J. (974): Variegated Pinangas. Principe*, 8:2224. HENDERSON, M.R. (959): Malayan Wild Flowers. Dicotyledons. Kuala Lumpur: Malayan Nature Society, Caxton Press. HIBBERT, S. (87): New and Rare BeautifulLeaved Plants. London: Bell and Daldy. KIEW, B.H. (compiler) (976): A Survey of the Proposed Sungai Danum National Park, Sabah. KIEW, R. (972): The Taxonomy and Ecology of Iguanura (Palmae). Ph.D Thesis, Cambridge University. (976): The Genus Iguanura (Palmae): Gdns* Bull Singapore 28: LEE, D.W. and LOWRY, J.B. (975): Physical Basis and Ecological Significance of Iridescence in Blue Plants. Nature (London) 254: 55. RICHARDS, P.W. (936): Ecological Observations on the Rain Forest of Mount Dulit, Sarawak. J. Ecol. 24: 37, (952): The Tropical Rain Forest. Cambridge: Cambridge University Press. RIDLEY, H.N. (923): The Flora of the Malay Peninsula. II. Gamopetalae. London. Reeve. SOEPADMO, E. and KIRA, T. (977): Contributions of the IBPPT Research Project to the Understanding of the Malaysian Forest Ecology. New Era in Malaysian Forestry STEENIS, VAN C.G.G.J. (969): Plant Speciation in Malesia, with Special Reference to the Theory of Nonadaptive Saltory Evolution. Biol. J. Linn. Soc. :9734. WALTER, H. (97): Ecology of Tropical and Subtropical Vegetation. (Trans. D. MuellerDombois). Edinburgh: Oliver and Boyd. {Received 23 August 978)

63 Pertanika, (2), 225 (978) Leg Paralysis Inducement by Risella 7 Oil in Lucilia sericata (Meig) as Affected by Sex, Time, Application Site, and oil Volume G. S. LIM Malaysian Agricultural Research and Development Institute Key words: Risella 7 Oil; Lucilia sericata (Meig); Leg Paralytic Induction, England. RINGKASAN Kejadian kaki lumpuh yang disebabkan oleh 'Risella7oil' he atas Lucilia sericata (Meig.) mempunyai kaitan yang bererti dengan jantina, masa y bahagian anggota yang dirawat dan isipadu 'oil' yang digunakan. Adalah diperhatikan bahawa ( Risella7oil' memberikan kesan yang lebih nyata ke atas lalat jantan dibandingkan dengan lalat betina. Walaubagaimanapun, bagi keduadua jenis lalat, sama ada jantan atau betina, kejadian kaki lumpuh bertambah apabila isipadu 'oil' digunakan dengan berlebihan PV$o (isipadu 'oil' yang diperlukan bagi menyebabkan tandatanda lumpuh ke atas 5% lalatlaiat yang diberi rawatan) ialah.3 W bagi lalatlalat jantan dan.2 pi bagi lalatlalat betina. Garisgaris regressi ialah y = 4. x +.57 bagi lalat jantan dan y = 2.64 x +.53 bagi lalat betina. Jumlah lalat jantan yang terkena kaki lumpuh adalah bergantung kepada bahagian anggota yang dirawat. Dengan menggunakan.3 pi 'oil', jumlah lalat yang terkena kaki lumpuh adalah dua kali ganda banyaknya bagi rawatan kepada abdomen dibandingkan dengan rawatan yang dilakukan ke atas thorax. Walaubagaimanapun tiada perbezaan yang bererti dapat dilihat di antara rawatan yang dibuat kepada bahagian belakang ataupun ke atas bahagian pedadaan abdomen. Lalatlalat yang menerima rawatan di bahagian kepala dan kaki tidak menunjukkan tandatanda lumpuh. Setakat ini, penemuanpenemuan yang diperolehi dari penyelidikan mencadangkan bahawa 'Risella7oil' boleh membawa lebih dnripada satu akibat. Namun begituy kepentingan dan peranannya dalam menyebabkan berlakunya lumpuh kaki serta diikuti dengan kematian adalah memerlukan penyiasatan yang lanjut dan lebih kritikal. SUMMARY Development of leg paralysis induced by Risella 7 oil on Lucilia sericata (Meig.) was found to be significantly affected by sex, time, application site, and oil volume. Male flies were noted to be more susceptible than female flies. In both sexes, more were affected at a given time when a greater volume of oil was applied. The PV S (volume of oil required to induce paralytic symptoms in 5% of the treated flies) was found to be.3 Mfor the male flies, and.2 &l for the female flies. Their probit regression lines are: y = 4.x +.57 for the male, and y => 2.64x \l 35 for the female. The number of male flies that developed leg paralysis was dependent on the site of application. With.3 ul of oil, there were about twice as many flies affected in treatment on the abdomen as compared to that on the thorax. However, no significant difference was observed between applications made dorsally and ventrally on the abdomen. Flies treated on the head and legs did not develop any paralytic symptoms. The findings suggest that there is more than one mode of action. However, the relative importance and role of any mode of action in contributing to the paralysis inducement with subsequent death, would require further and more critical investigations. INTRODUCTION including varying dosages applied topically, by injections, or as surface residues (Bard, 96; Risella 7 oil is commonly used in insecticide Lewis, 962, 963; Busvine, 962, 97). formulations. In blowfly studies, it has been Although no inducement of leg paralysis was used alone or with insecticides; the treatments noted in these investigations, the oil was later Taken in part from a thesis for the D.I.C and M.Sc degree of the University of London. The study was conducted at the Field Station, Silwood Park (Berkshire), Imperial College of Science and Technology, London. 2

64 G. S. LIM found to induce leg paralysis with the blowfly, Lucilia sericata (Meig.) (Lim, 972, 976). Further studies were conducted to investigate the effect of the oil, particularly on the pattern of paralytic development (Lim, in press). However, the present investigations on paralytic inducement as affected by sex, time, application site, and oil volume, were aimed at providing deeper insights into their nature and relationships, well as a better understanding of some aspects on the mortality process since the paralysis is noted to be an extended state of moribund condition (Lim, 972; 976). MATERIALS AND METHODS Throughout the studies, the temperature was maintained at 26 ^ C and the relative humidity 65 ± 5%. All the flies used were unmated and of the same age (4 days old) and brood. Within 24 hours of emergence, the males were segregated on the basis of the distance between the eyes, which in the female is approximately more than onethird the total width of the head (Aubertin, 933). This early separation enabled the flies to remain unmated and their age known throughout the studies. During treatment the flies were temporarily immobilized by chilling. Application was made with a footoperated Burkard microapplicator fitted with an ordinary ml glass syringe that carried a bent needle with a blunt tip. Each fly was topically applied with a nominal dose of.3 as delivered by the applicator. Except for flies used in studies concerning sites of application, the treatment was on the anterior margin of the second last abdominal segment and on the ventral surface. Flies that were deformed or crippled by the necessary handling were destroyed and replaced. from application (Fig. ). For a particular volume of the oil applied, the number affected increased with time. Depending on the volume used and the sex of the treated flies, the effect may be exhibited as early as two hours after treatment. From then on, the number affected increased rapidly; the maximum (or limiting) number was reached by the second or third day after treatment when male flies were topically dosed with.2 and.3 /AI of oil. No significant increase in the number of affected flies was observed after this time. Any fly that did not show any symptom then remained unaffected. 2. Fly number affected in relation to sex The male and female flies showed differing susceptibility to the paralytic effects induced by Risella 7 oil. With a given volume, the male was observed to be more susceptible, generally having a higher maximum number of affected individuals (Fig. ). Also, the time taken to reach the limiting level was faster. This was probably related to the different body weight and size of the two sexes, the male being much smaller (mean weight ± S.E. = 2.64 ±.99 mg) than the female (mean weight ± S.E. = 3.72 ±.7 mg). 3. Fly number affected in relation to the volume applied The maximum number of flies affected was noted to depend on the volume of Risella 7 oil applied. In general, it increased with higher volume. For instance, with.2 /xl and. ^ of oil, the male flies showed a maximum of about 9% and 36% affected, respectively (Fig. ). This general relationship was also observed in the female flies, although the rate of development and maximum number affected were comparatively lower for any given volume studied. After treatment, the flies were kept in round plastic containers ( cm diameter x 3.7 cm high) in groups of ten flies/container. Water and food in the form of granulated sugar were provided. At regular intervals, the flies were observed for symptoms of paralysis; each fly was considered paralytic when it had at least two legs paralysed. For each set of study, a parallel group of untreated flies was used as control check. RESULTS. Fly number affected in relation to time from application The numbtr of flies affected by Risella 7 oil was observed to vary with the time interval 2 The volume of oil required to induce paralytic symptoms in 5% of the treated flies (PV 5 ) was determined for both the sexes. The regression lines obtained (Fig. 2) showed that in both the sexes, a linear relationship existed between the volume of the oil applied (log x) and the percentage suffering leg paralysis (probit y). Their regression equation are: y = 4.x +.57 for the male, and y = 2.64 X +.53 for the female. In the latter, the PV 5 value is.2 Ml with 95% fiducial limitsof.il M to.37^. This is almost double that of the male which has a PV5 value of.3 /A with 95% fiducial limits of.8 /xl to.9 fx\. Evidently, the difference is related to the body weight since the PV 5 for both sexes are almost the same when expressed against the weight,.6 /xl/mg for the male and.66 /xl/mg for the female.

65 LEG PARALYSIS INDUCEMENT BY DISELLA 7 OIL IN LUCILIA SERICATA r o 3 ul f 2 ul uf HOURS DAYS Period after treatment Figure. Number of Lucilia sericata developing leg paralysis in relation to the volume of Risella 7 oil applied, time interval from application, and the sex of the flies. (Number of test flies/treatment: male = 3 4, female = 4 4) 4. Fly number affected in relation to the site of application In this study, male flies were randomly divided into 5 treatment groups. One of the groups was used as an untreated control while the remaining flies were treated topically with.3 //.I Risella 7 oil according to their treatment groups which were as follows: dorsaliy on the head, dorsally on the thorax, dorsally on the abdomen, and ventrally on the abdomen. The results showed that flies treated on the abdomen were about twice as susceptible than those which had applications on the thorax (Fig. 3). Among those treated on the abdomen, no significant difference was observed between those treated dorsally or ventrally. Throughout the study, neither control flies nor flies treated on the head appeared to be affected. 5. Risella 7 oil treatment and fly mortality This study on the toxic effect of Risella 7 oil (applied at.3 /el) in relation to insect mortality, is based on the pooled data obtained from the 22 various experiments conducted. A total of 2 treated male flies and 258 untreated male flies were studied. As noted in previous studies, leg paralysis in treated flies occurred within two hours of treatment, increasing rapidly with time as was noted in more than 8% of the flies by two days following treatment. This pattern of development was observed, irrespective of whether the percentage paralysis calculated was based on only the surviving individuals, or with the dead individuals included under paralysis (Fig. 4). In the study, a significant number of treated flies was observed to die following paralytic symptoms. This mortality was also noted to increase with time, reaching 66% at 7 days after treatment as compared to 3% in the untreated control (Fig. 4). Mortality began to set in around 24 hours after treatment. The significantly higher mortality observed suggested that the applied oil, under the conditions studied, had some toxic effect on the insect.

66 G. S. LM PV, MALE(o)= I FEMALE i )= QD O *4 LOG x CONCENTRATION (ul) 6 Figure 2. Regression lines for the calculation of the PVSQ (volume of Risella 7 oil which induced leg paralysis in 5% of the treated flies), for both male and female Lucilia sericata (Meig.). The oil was applied topically. (Number of test flies/concentration: male == 34, female == 44). DISCUSSION From the investigations, it was evident that the number of flies affected by Risella 7 oil is closely associated with the time from exposure, the volume of oil to which flies were exposed, the site of application, and the sex of the flies. The males were noted to be more susceptible; in both 23 sexes generally, the effect became more pronounced with increasing volume of oil applied. At high volume, the oil showed toxic effect. Presently, the mode of action of the oil in inducing leg paralysis and subsequent death (for some) is still unclear. The findings suggested that the oil could act through the spiracles by

67 LEG PARALYSIS INDUCEMENT BY DSELLA 7 OIL IN LUCILIA SERICATA A a Thorax _ 8 3 * Abdomen (Dorsal ) Abdomen (Ventral) B Treatment on Head ) ) i =None affected Untreated Control ) 2 4 HOURS Period after treatment 4 5 DAYS Figure 3. Number of Lucilia sericata (Meig.) developing leg paralysis in relation to the site of application of the Risella 7 oil. (Treatment: topically with.3 JUL/; Number of test flies/treatment: 3 35 males.) ^ ^T.~. Z.".* Paralysis (Deads included) 8 Paralysis (Dead Excluded) I 6, Mortality (Treated) o 4 2 Mortality (Control) 2 4 HOURS I DAYS Period after treatment Figure 4. Effect of Risella oil, topically applied at.3 Vllfly, on the leg paralysis and morality of the male Lucilia sericata (Meig.) 24

68 either physically blocking them and affecting the oxygen supply, or by entering rapidly into the body and to the vital organs through the spiracles and tracheal system. Whichever the route the mechanism of the paralytic induction is basically related to the nervous system. From the present studies, it was found that entry through the spiracles had special significance since flies treated on the head, where there were no spiracles, did not exhibit any paralytic symptoms; and thoracic treatments showed less effect than treatments on the abdomen which has many more pairs of spiracles. Alternatively, the oil may dissolve and penetrate directly into the body through the cuticular layer. Such a mode of entry by chemicals has been reported or implied in reviews on this subject by several workers (Brown, 95; Richards, 95; 953; O'Brien, 967; Wigglesworth, 942; 948; and Ebeling, 964). Based on this concept, the greater effect noted in the abdominal over the thoracic treatment may be explained by the greater surface area in the former. This allows the oil, which spreads readily on application, to act more effectively. However, since no paralytic symptoms were exhibited when treatment was made on the head, it would appear that the vital site of entry which resulted in leg paralysis, does not lie in the head. Otherwise, any direct penetration of the oil through the head cuticle should eventually also result in leg paralysis, unless differential cuticular penetration exists (Lewis, 965), with little or no penetration taking place through the head. The possibility of the leg paralysis being induced by the oil from purely physical effects on the legs directly was checked by treating flies on the legs only Flies were treated with.3 //.I of oil, either on the forelegs, midlegs, or hindlegs. In this study, the possibility that some of the oil applied on the legs may eventually creep up onto the body (Lewis, 962) and induce leg paralysis in a manner similar to treatments that were made directly on the thorax or abdomen was considered. However, since no leg paralysis developed with such treatment, the cause of leg paralysis purely from the physical effects of the oil on the legs would thus appear to be unlikely. The findings of this study suggest that there is more than one mode of action. However, the relative importance and role of any one mode of action in contributing to the paralysis inducement with subsequent death, would require further and more critical investigations. ACKNOWLEDGEMENT Grateful thanks are due to my supervisor, Dr. C. T. Lewis, for his constant interest and guidance. G. S. LIM 25 REFERENCES. AUBERTIN, D. (933): Revision of the genus Luciha RD. (Diptera, Calliphoridae). J. Linn. Soc Zoology, 38: BARD, J.M. (96): Influence of the solvent on the toxicity of injected solutions of an insecticide. M.Sc. Thesis, University of London. 3. BROWN, A.W.A. (95): Insect control by chemicals. London: Chapman and Hall. 4. BUSVINE, J.R. (962): A laboratory technique for measuring the susceptibility of houseflies and blowflies to insecticides. Laboratory Practice: BUSVINE, J.R. (97): A critical review of the techniques for testing insecticides. Commonwealth Agr. Bureaux, Slough. 6. EBELING, W. (964): "Permeability of insect cuticle." The physiology of insecta. Rockstein, M. (Ed), pp , Academic Press, New York. 7. LEWIS, C.T. (962): Diffusion of oil films over insects. Nature. 93: LEWIS, C.T. (963): Some applications of radioisotopes to the study of the contamination of insects by insecticide solutions. Proc Ser. Int. atom. Energy Ag. 74: LEWIS, C.T. (965): Influence of cuticle structure and hypodermal cells on DDT absorption by Phormia terraenovae RD. J. Insect PhysioL, : LIM, G.S. (972): The effect of chemicals applied topically on Lucilia sericata (Meig.) with special reference to Risella 7 oil. M.Sc. Thesis. University of London.. LIM, G.S. (976): Inducement of leg paralysis in blowfly {Lucilia sericata Meig.) by Risella 7 oil. Mai Agric. Res., 5: LIM, G.S. (in press): Notes on the developmental pattern of leg paralysis induced by Risella 7 oil on Lucilia sericata (Meig.). The Planters Guild Bulletin. 3. O'BRIEN, R.D. (967): Insecticides: action and metabolism. New York: Academic Press. 4. RICHARDS, A.G. (95): The integument of arthropods. Minneapolis: University of Minnesota Press. 5. RICHARDS, A.G. (953): "The penetration of substances through the cuticle." Insect physiology. Roeder, K.D. (Ed.), pp London: Chapman and Hall. 6. WIGGLESWORTH, V.B. (942): Some notes on the integument of insects in relation to the entry of contact insecticides. Bull. ent. Res. 33: WIGGLESWORTH, V.B (948): The insect cuticle. Biol. Rev. 23:4845. (Received 6 August 978)

69 Pertanikal (2), 2635 (978) Progress of Natural Regeneration after Final Felling under the Current Silvicultural Practices in Matang Mangrove Reserve Key Words: Mangrove, Natural Regeneration P. B. L. SRIVASTAVA and DAUD KHAMIS Faculty of Forestry, Universiti Pertanian Malaysia RINGKASAN Kertas ini membincangkan keputusan penyelidikan he atas kemajuan pemulihan hutanhutan bakau selepas tebangan, khususnya susunan jenis, pola taburan dan kepadatan stok. Penyelidikan tersebut telah dibuat di Matang Hutansimpanan Bakau dalam Renj Port Weld. Kawasankawasan yang dipilih adalah dari kelaskelas 'inundation' III dan IV (Watson, 928) dan mengandungi tumbuhan jenis Rhizophora yang biasa. Pemulihan di dua kelas umur, iaitu 2 bulan dan 24 bulan selepas tebangan, telah dibanci secara Bancian Linear (LRS). Rhizophora spp. dan Bruquiere parviflora adalah jenisjenis yang penting terdapat di diriandirian baki. Bilangan anakbenihanakbenih Rhizophora spp. di kawasankazvasan 2 dan 24 bulan selepas tebangan adalah 37 dan 3376 seekar. Bilangan anakbenihanakbenih B. parviflora pula di kawasankazvasan 24 bulan selepas tebangan merosot secara berkesan, kepada 62 seekar jika dibandingkan dengan 282 anakbenihanakbenih seekar di kawasan 2 bulan selepas tebangan. Anakbenihanakbenih Rhizophora spp. tidak mati berluasan di dalam jangkamasa tersebut. Kawasankawasan di keduadua kelas umur mempunyai bilangan anakbenihanakbenih yang mencukupi untuk mewujudkan dirian yang bercukupan stok pada hujung giliran sekiranya kematian yang berluasan tidak berlaku. SUMMARY The paper presents the results of an investigation on the progress of regeneration of mangrove forests after logging with emphasis on species composition, distribution pattern and stocking. The investigation was undertaken in the Port Weld Range of The Matang Mangrove Reserve. The areas chosen belong to inundation classes III and IV (Watson, 928) representing a typical Rhizophora type. Two age classes, i.e. 2 months and 24 months after felling, were sampled by means of Linear Regeneration Sampling (LRS ) method. Rhizophora spp. and Bruguiere parviflora were the most important species in the residual stands with 37 and 3376 seedlings per acre of Rhizophora spp. respectively in the areas 2 and 24 month after final felling. There was a significant decrease in the number of B. parviflora seedlings in the areas 24 months after felling with only 62 seedlings per acre compared to 282 seedlings per acre, 2 months after felling. There was no large scale mortality in the Rhizophora seedlings with time during this period. At both the age classes, the areas had enough number of seedlings to produce a fully stocked stand at the end of rotation if no large scale mortality occurred. INTRODUCTION though there was heavy damage to the existing In a study conducted simultaneously, Sani seedlin S cr P durin g W^ng and transporting (977) discussed the effect of logging at the time operations, there was still a sufficient number of of final felling on different aspects of natural seedlings in the residual stands to give an anticiregeneration in a typical Rhizophora type in pated yield provided no mortality due to exposure Matang Mangrove Reserve. It was found that or any other cause occurred. Saham Pahang Berhad, Kuantan 26

70 This study reports the progress of natural regeneration during the two years after felling in the same locality and type of forest. So far, there has been no quantitative information on the progress of natural regeneration of Rhizophora spp. In many areas, natural regeneration is deficient due to yet unknown causes and the State Forest Department has to resort to planting to obtain adequate stocking for sustained yield. Systematic studies are, therefore, needed to determine the progress of natural regeneration of Rhizophora species after the final felling in some typical localities. REVIEW OF WORK Watson (928), writing at a time when the current techniques of treating a mangrove stand were only just beginning to be tried out, provides a comprehensive account of these forests which is true to this day. Noakes (95, 952) observed that (i) no systematic investigation into the problem of regeneration had ever been attempted, perhaps because the treatment of mangrove forest had gradually come to be regarded as standardized; and as natural regeneration was expected to succeed over twothird of the annual coupe, planting was not unduly expensive and the rotation was long enough to "even off" the differences, (ii) productive ability of Rhizophora spp. was high; fruiting began at four years of age and though the number of seed varied considerably from year to year, there was sufficient seed available to restock all felling areas within two or three years, (iii) Achrostichium aureum (Piai) which occurred throughout the whole range of Rhizophora forest responded readily to full light and acted as a nurse to the existing seedlings; and if there were no seedlings then few could get in through tidal waves. Such areas could only be stocked by retaining seed bearers, (iv) the adverse influence of slash on the regeneration could be minimised by constant vigilance to ensure that all utilizable wood was removed from the felling area, and by lowering the rotation age thereby reducing the crown size of the final crop, (v) water could play an effective role in the dispersal of seeds of Rhizophora spp. and in natural regeneration only after the slash and most of the stilt roots of the felled trees had decayed, and (vi) large areas could not be regenerated with coppice seedlings or artificially. Dixon (959) questioned the role of the waterborne fruits in the normal process of natural regeneration of Rhizophora forest and suggested that the continuous disturbance by subsequent P. B. L SRIVASTAVA AND DAUD KHAMIS 27 tides was the main cause for poor regeneration; he also pointed out that though the earlier workers had been aware that young seedlings of Rhizophora damaged by logging could recover by coppice shooting, there was no information available as to the number of seedlings capable of coppicing after the final felling; he maintained that while the larger blanks were filled up by planting two years after exploitation, the smaller blanks were filled up naturally after about seven years, provided that weeds such as Derris uliginosa and Achrostichium aureum did not choke the site. The situation has changed much during the last two to three decades. Firstly, with the rise in population and increased demand for the mangrove wood, the area of exploitation has inadvertently been extended which will require more.effort to ensure natural regeneration in the residual stands. This applies to the Peninsula as well to Sabah and Sarawak. Secondly, the high cost of planting and shortage of staff at all levels to regenerate the deficient areas in the stipulated period are likely to restrict future planting. Thirdly, adequate quantities of seed may not be available for planting. Fourthly, large areas of forest have been classified as unproductive in Matang Mangrove Reserve on the grounds that the crop consists of Bruguiera spp. Dixon (959) pointed out that areas now due for felling carry a much greater proportion of this species in the crop. According to Mohd. Darus (969) more than 5 acres of potential Rhizophora forest in Matang Reserve were degraded on account of the presence of B. parviflora. Paul Chai (974, 975) draws attention to many problems faced in regenerating the exploited mangrove forests of Sarawak: mounds, wherever they exist, are almost immediately invaded by A. aureum which become particularly abundant in the exploited areas; the soil in the area is greyish brown, thick and sticky clay and is nonporous and insufficiently aerated; regeneration of Rhizophora spp. and Bruguiera spp. on this type of soil was found to be very scarce except on the soft mud near river and stream banks. The seedlings do not get anchored easily, and even if they do, most of them fail to develop. The Sarawak Forestry Department plans to study the enrichment planting of loggedover areas as a means of regenerating the mangrove forests and removal of slash in experimental logging blocks to ascertain the possibility of improving the regeneration environment. Abdul Manap and Srivastava (975) have suggested that studies be carried out to ascertain the rate of decomposition of slash after logging and its effect on natural regeneration.

71 NATURAL REGENERATION AFTER FINAL FELLING IN MATANG MANGROVE Yusuf (977) suggested that there is a need for more intensive studies before logging is extended over large mangrove areas because the swampy habitat is also believed to be an important nursery for many species of marine fish and prawns. Recent studies in Sabah have shown that 82 per cent of the prawns exported from Sabah were of a species that spends a greater part of its lfe cycle in swamps and are caught in the coastal waters close to swamps. Silviculture The silviculture of the mangrove forests is fairly simple compared to that of the inland forests because the former are more homogenous and uniform. A clear felling system is used. All useless species left behind after harvest are slashed or poisoned. In many workedover stands, blanks form about 253 per cent of thr harvested area. These areas are planted with Rhizophora spp. usually two years after logging when most of the slash has decomposed. In the Achrostichium aureum dominated areas planting is recommended immediately after logging. The seeds of R. mucronata are planted at 6' x 6' and those of R. apiculata are spaced at 4' X 4'. The planting operation is normally carried out during the months of July November to coincide with the fruiting season of Rhizophora spp. There are three thinnings, all mechanical, in a 3year rotation cycle: I. thinning at the age of 5 years with 4' stick; II. thinning at the age of 92 years with 6' stick and III. thinning at the age of 2526 years with 7' stick. The current silviculture prescriptions are described by Mohd. Darus(969). METHOD OF STUDY (i) Description of the experimental area This investigation was undertaken in the Matang Mangrove Reserve. All the study plots are situated in the Port Weld range (Table ). The areas chosen for the present study belong to inundation classes III and IV (Watson, 928) representing a typical Rhizophora type which forms the greater part of these swamps. Seventy to ninety per cent of the growing stock is dominated by Rhizophora species, R. apiculata and R. mucronata. The other inundation classes were not covered in the present study because they could be too wet or too dry and would have introduced an additional factor which affects natural regeneration. These sites were selected for detailed sampling because they were typical sites where no factor had apparently adversely affected the progress of natural regeneration. However, a clear differentiation between the two inundation classes is not apparent particularly in regard to the quality of crop and the regeneration environment. (ii) Field procedures In the collection of data on the composition, distribution, and progress of regeneration and percentage survival of the seedling crop of two age classes, i.e. 2 months and 24 months after logging, Linear Regeneration Sampling procedure (LRS ) (Anon, 975) was adopted. This method has been standardized for the assessment of natural regeneration in the inland dipterocarp forests and was successfully used by Liew et al. (977) in Sabah mangroves. A base line was drawn along a convenient boundary, such as a channel or compartment limits, and sampling lines were laid out on this base line at intervals of chains. Milliacre quadrats were laid out contiguous to one another on the right side of each sampling line. The occurrence of seedlings of different species was recorded in the milliacre quadrats by complete count in three height classes, viz., Ho ('), H, (l'5') and H 5 (5'). RESULTS AND DISCUSSION () Composition of seedling crop As many as nine species were recorded in areas 2 months after felling, viz., Rhizophora apiculata, R. mucronata, Bruguiera parviflora, B. gymnorrhiza, B. sexangula, B. cylindrica, Avicennia species, Exoecaria agallochia and Heritiera littoralis. In areas 24 months after felling, the latter three species were not encountered. However, at both stages, Rhizophora species were the dominant constituents and B. parviflora was the most important associate in terms of frequency, density and abundance. In areas 2 months after felling, Rhizophora species seedlings constituted over 7 per cent of the crop (except in plot 3) and together with B. parviflora, made up over 99.9 per cent of the seedlings. In areas 24 months after felling, the proportion of B. parviflora was further reduced. The crop became more pure with over 96 per cent of the seedlings belonging to Rhizophora species, mainly R. apiculata (Table 2, Fig. and 2). /?. mucronata was present in the soft mud along the banks of the rivers and channels. The discussion that follows deals with different aspects of the two most important species. 28 (2) Abundance of natural regeneration A wide variation was noted in the density of Rhizophora species in different plots, 2 months after felling (Table 3). It was possible to deter

72 P. B. L SRIVASTAVA AND DAUD KHAMIS TABLE Sampled sites and date of sampling Stage Date of Sampling Compt. No. Plot No. Coupe No. Area (acre) No. of milliacre quadrats. 2 months after felling (A) / (B) 2 53/ (A) 3 27/ (B) 4 3/ Total Acreage months after felling /74 35/73 3/ Total Acreage. TABLE 2 Percentage Stocking by Species Stage Plot No. Rhizophora spp. B. parviflora Rhizbphora spp. & B. parviflora B. parviflora & other spp. Other spp. 2 months after felling Av months after felling Av TABLE 3 Abundance of seedlings in different plots, 2 months after felling 95% Confidence limit Plot No Mean/acre Standard Error S.E.% Lower Confidence limit per acre Upper Confidence limit per acre Number of plants per acre Rhizophora spp. B. parviflora

73 NATURAL REGENERATION AFTER FINAL FELLING IN MATANG MANGROVE < CL a O T3 csi o _O CL L_ o _c a O N *_C cr CL gui( m LO u O Q. t/ o o o CO o CM 3

74 P. B. L. SRIVASTAVA AND DAUD KHAMIS I (/) g> 8 6 C O I 2 plot 5 plot 6 plot 7 Average Localities Rhizophora Bruguiera species parviflora Figure 2. Histogram showing percentage stocking by species for areas 24 months after felling. other species mine the coefficient of variation for a complete count of the seedlings was carried out. The maximum number of seedlings (547/acre) was recorded in plot and the minimum (647/ acre) in plot 3. The variation in the case of B. parviflora was greater (c.v. = 59 per cent) than in Rhizophora species (c.v. = 42 per cent). Rhizophora seedlings outnumbered B. parviflora by nearly three times. There were, on an average, 37 seedlings of Rhizophora species per acre with a standard error of 2 per cent and a range of seedlings per acre compared to 282 seedlings per acre of B. parviflora with a range of per acre with a standard error of 29 per cent (P<.5). 3 At this stage the most abundant height class in both species was Hi (the seedlings between one to five feet), accounting for 9 per cent of the seedlings of Rhizophora species and 85 per cent of B. parviflora. The other two classes were poorly represented, H o having the least number of seedlings in Rhizophora species and H 5 in B. parviflora. The number of seedlings of B. parviflora was four times that of Rhizophora species in H o class. But in H { and H 5 height classes, Rhizophora species seedlings were three and seven times more abundant than B. parviflora respectively. These two characteristics i.e. predominance of Rhizophora species and B. parviflora and Hj height class appear to be typical of this

75 NATURAL REGENERATION AFTER FINAL FELLING IN MATANG MANGROVE age group. The two species made up 9 per magnitude of variation in the number of seedlings cent of the residual stand. of the two most dominant species i.e. Rhizophora species (c.v. = 24 per cent) and B. parviflora The analyses of different aspects of density (c.v. = 36 per cent) was reduced. In the case of seedlings in areas 24 months after felling are of R. apiculata, the number of seedlings varied shown in Tables 5 and 6. It is evident that the between per acre in different plots Plot No Av. o< / O Ho 3 TABLE 4 Number of seedlings per acre in different height classes, 2 months after felling Rhizophora spp Hi Hs Ho B. parviflora Hi H Ho Total Hi H Grand Total Plot No. TABLE 5 Abundance of seedlings in different plots, 24 months after felling Rhizophora spp. Number of plants per acre B. parviflora Mean/acre Standard Error S.E. % 4 2 Lower confidence limit per acre % Confidence limit Upper confidence limit per acre r>i A. Plot No Av. o /o Ho 32 TABLE 6 Number of seedlings per acre in different height classes, 24 months after felling 6 3 Rhizophora spp. Hi H Ho B. parviflora Hi H Ho Total Hi H Total

76 with an average of 3376 seedlings per acre and a range (P<.5) and a standard error of 4 per cent. In the case of B. parviflora, the number varied between 37 8 per acre in different plots with a mean of about 62 seedlings per acre and a range of 68 (P<.5) and a standard error of 2 per cent. At this stage Rhizophora species seedlings were even more abundant, being about 5 times as many as those of B. parviflora seedlings. Hj was still the predominant height class though the number of seedlings of B. parviflora was slightly more in H 5 than in H]. The proportion in H] seedlings, however, decreased substantially from 9 per cent to 57 per cent in Rhizophora species and from 85 per cent to 37 per cent in B. parviflora within a year. H o class was very poorly represented. In all cases Rhizophora seedlings were more abundant than B. parviflora, Ho (3 times), Hi (8 times), H 5 (nearly 5 times). This indicates that there was almost no addition to the natural regeneration in the form of new recruitment during the two years after felling; the present crop of seedlings, which shows progressive height growth, is the one which survived the onslaught of the final felling. (3) Mortality In general, there was a decrease in the number of seedlings with time after logging. As against an average of 37 seedlings per acre of Rhizophora species 2 months after felling, there were 3376 seedlings per acre of these species 24 months after felling. A much higher rate of mortality was observed in the seedling crop of B. parviflora. There were, on average, only 62 seedlings per acre 24 months after felling compared to 282 seedlings per acre 2 months after felling (Tables 3,5). While a low rate of mortality in the seedlings crop of Rhizophora may be due to "natural thinning" of the excess number of seedlings in a few overcrowded patches, the cause for the sharp drop in the number of B. parviflora seedlings are not easy to find. It may be due to both interand intraspecific competition. Almost all species found in the swamp are light demanders and therefore the high rate of mortality of B. parviflora seedlings in these areas may be due to shade created by the more vigorously growing Rhizophora seedlings. The analysis also showed that most of the dead seedlings of B. parviflora were smaller, belonging to height classes less than five feet. The death of the young seedlings could also be due to heat and dehydration brought about by the decomposition and collapse of stilt roots, stumps and other remains of the felled trees with time,* thus depriving the young seedlings of P. B. L. SRIVASTAVA AND DAUD KHAMIS 33 shade. The movement of partially decomposed slash during flooding can be another cause. In fact, Watson (928) considered movement of semidecomposed slash due to tidal action one of the most important causes of mortality of tender seedlings which accounts for blanks in the loggedover areas. The damage caused by the attack of crabs or monkeys (Noakes, 95) appears to be negligible in the naturally regenerated stands at this stage. Liew et at (977) also noted large scale mortality of B. parviflora seedlings with time. (4) Adequacy of natural regeneration (i) 2 months after felling In the inland forests, an area is considered adequately regenerated naturally if it has a minimum stocking of 3 well distributed seedlings of important timber species per acre with an allowance for 2 per cent mortality (Anon, 975). However, no such criterion exists for any species of mangrove swamps in the Peninsula. The figure acceptable for inland forests would not apply to the mangrove swamp species because the growth rates of the species and the silviculture system are different. In the residual stands of the mangrove forests, even if each milliacre plot contains a minimum of one seedling, the stocking would be only seedlings per acre which is far below the desired stocking needed for the development of a productive stand. It is assumed (Sani, 977) that the number of stems present at the time of final felling varies between 5 8 per acre and planting with Rhizophora apiculata is carried out at 4' X 4' yielding 2722 seedlings per acre. The number of stems per acre after the first, second and third thinning will be 2722, 2, 889, respectively. These computations take into account normal rate of mortality, if any, so as to yield a well stocked stand at the time of final felling. This means that there should be a minimum of three seedlings per milliacre plot in the regenerated stands before the first thinning is carried out if a potentially productive stand for the next rotation is to be developed. On the basis of the above assumption, for an area to be adequately stocked, there must be at least per cent of the total milliacre plots with fvwt seedlings each, or 68.6 per cent containing at least four seedlings per plot, or 9.75 per cent with at least three seedlings per plot in order to have at least 2722 stems per acre at the time of first thinning. The average number of the seedlings (irrespective of height) of the preferred species (Rhizo

77 NATURAL REGENERATION AFTER FINAL FELLING IN MATANG MANGROVE phora spp.) per acre is more than 2722 except in plot 3 which carried only 647 seedlings per acre (Table 3). The analysis of the basic data further indicates that the seedlings are evenly dispersed in all the stands sampled in the present study. As many as 6 per cent of the plots had two or more seedlings and the blanks constituted only about 25 per cent of the plots. This is probably due to the fact that the crop on the most productive sites in this area is homogenous, mainly of Rhizophora species. Plot 3 (Tables 3 and 4) shows an interesting distribution pattern of Rhizophora species and B. parviflora seedlings. The site seems to favour the growth of the latter. By studying such sites in detail it is possible to identify the factors responsible for the emergence of this 'weed' species on a large scale in a typical and potential Rhizophora stand after final felling. (ii) 24 months after felling At this age also, as is evident from the tables 5 and 6, plots 6 and 7 have more than an adequate number of seedlings of the preferred species on the same criteria. Only plot 5 is slightly understocked. However, the percentage of blank plots is increased slightly indicating that there has been no new recruitment during this period either through waterborne seeds or from the seed bearers. It was surprising to note that there was a significant decrease in the number of B. parviflora seedlings during this period. It comprised only 2.5 per cent of the total area. This appears contradictory to the observations of workers who had noted that this species tended to increase in number with age in the regenerated stands. It is probable that the incidence of B. parviflora after felling is greater in other inundation classes, such as V and VI, though it may dominate even potential Rhizophora stands, as in the case of plot 3 (Tables 3 and 4). Similar observations were made by Dixon (959) and Mohd. Darus (969). It is also possible that B. parviflora might start appearing three or four year after felling, ultimately forming an important associate of Rhizophora species. Further studies of more age classes, at least up to the stage of I thinning, and in the remaining inundation classes, need to be undertaken to determine the factors responsible for the predominance of B. parviflora in a typical Rhizophora type. It was further noted that the number of Rhizophora seedlings of Hi class had gone up to 463 per acre (Table 6) as against only 32 seedlings per acre (Tables 4) during the period of one year. This indicates the rate of growth of this species under favourable conditions. On the other hand there was a sharp drop in Hj 34 height class of B. parviflora seedlings from 95 sedlings per acre (Table 4) at 2 months after felling to only 23 seedlings per acre (Table 6) at the end of 24 months. The probable reasons for this decrease have already been stated. CONCLUSIONS On the basis of the present study in a Rhizophora species dominated forest on inundation classes III and IV, the following conclusions can be drawn:. Rhizophora spp. and B. parviflora are the most important species in the residual stands at 2 and 24 months after felling. Seedlings of other species are rarely present. 2. There is no large scale mortality in the Rhizophora seedlings with time. 3. A large number of B. parviflora seedlings died between 2 and 24 months after final felling probably because of severe competition from Rhizophora seedlings, or exposure and dehydration, or both. 4. Most of the stands sampled in the present study were found to be adequately stocked, 2 and 24 months after felling. ACKNOWLEDGEMENTS The authors are indebted to the Director of the Perak State Forest Department and Encik Xik Answar bin Nik Salleh (A.D.F.O. Taiping) for permission and help in various forms to carry out the survey. Thanks are also due to Enchik Zainuddin, En. Khairi, and En. Majid, members of the staff of Port Weld Range, for their assistance. Special thanks are due to Professor Haji Abdul Manap Ahmad, the Dean Faculty of Forestry, Universiti Pertanian Malaysia, for approval of the project, and for his encouragement, critical suggestions, and for providing the field staff (Encik Harun, Encik Rahim, Encik Haidzir and Encik Ahmad). The authors are also thankful to Dr. G.M. Bonnor and Mr. M.N. Asthana for their help in the data analysis. Lastly, one of us (DK) wishes to thank his friend Abdullah Sani and his family for their hospitality during the period of his stay at Taiping. REFERENCES ABDUL MANAP AHMAD and SRIVASTAVA, P.B.L. (975): Effects of logging in mangrove vegetation in Peninsular Malaysia. ILP.M. (Unpublished).

78 P. B. L. SRIVASTAVA AND DAUD KHAMIS ABDULLAH SANI BIN SHAFFIE. (977): Effect of logging on the natural regeneration of Rhizophora species under the current silvicultural practices in Matang Mangrove Reserve. B.S. Thesis (Unpublished), Universiti Pertanian Malaysia. ANON. (975): A guide to Linear Regeneration Sampling One (LRS ). Pamphlets, Forest Department, Malaysia. DIXON, R.G. (959): A working plan for the Matang Mangrove Forest Reserve, Perak. Forest Department Publication. LIEW, T.C, DIAH. MOHD. NOR, and WONG, Y.C. (977): "Mangrove exploitation and Regeneration in Sabah." A New Era in Malaysian Forestry. Kuala Lumpur (Ed. Sastry, Srivastava and Manap), Universiti Pertanian Malaysia. MOHD. DARUS HJ. MAHMUD. (969): Rancangan Kerja bagi hutan simpanan paya laut Matang Perak. Forest Department Publication. NOAKES, D.S.P. (95): Notes on the Silviculture of the Mangrove Forests of Matang, Perak, Malayan For. 4(4): (952): A working plan for the Matang Mangrove Forests, Perak. Forest Department. Federation of Malaya Publication. PAUL, P.K. CHAI. (974): The potential of Mangrove Forests in Sarawak. Malayan For, 37(4): (975): Mangrove Forest of Sarawak. Malayan For. 38(2): 8. WATSON, J.G. (928): Mangrove Forests of the Malay Peninsula. Malayan For. Rec (6). YUSUF NAIR. (977): An appraisal of the economic potential of Mangrove swamps, M.S. Thesis (Unpublished), Universiti Pertanian Malaysia. (Received 28 August 978) 35

79 Pertanika (2), 3642 (978) Mineralogy and Related Properties of Acid Sulphate Soil from Melaka, Peninsular Malaysia SHAMSUDDIN JUSOP and TUAN OMAR RAJA SENIK Department of Soil Science, Faculty of Agriculture, Universiti Pertanian Malaysia Key words: Acid Sulphate, Montmorillonite, Pyrite RINGKASAN Sifat mineralogi, kimia dan fizik tanah asid sulfat dari Kuala Linggi, Melaka telah dikaji. Tanah ini didapati mengandungi mineral montmorillonit, kaolinit, illit, klorit dan pirit. Tidak ada perbezaan dari segi mineralogi diantara lapisan atas dengan lapisan bawah, tetapi kuantiti montmorillonit dan pirit mungkin bertambah ke bawah. Tanah asid sulfat ini mungkin berkeupayaan untuk menghasilkan pengeluaran pertanian yang tinggi; kehadiran montmorillonit meninggikan keupayaan pertukaran kation (melebehi 6 m.e./loog tanah). Walau bagaimana pun, kehadiran pirit yang senang teroksid kepada asid sulfat apabila air disalirkan, menyebabkan potensil pertanian tanah ini rendah. jfumlah sulfat bertambah ke bawah, begitu dengan sulfat larutair. Sekiranya pirit itu dapat dimusnahkan dengan tidak menjejaskan sifatsifat fizik dan kimia mineral lain, mungkin tanah ini mempunyai potensil pertanian yang tinggi. SUMMARY Miner a logical, chemical and physical properties of acid sulphate soil from Kuala Linggi, Melaka, were studied. The study shows that the soil contains kaolinite, montmorillonite, illite, chlorite and pyrite. There are no mineralogical differences between the top and the lower horizons. Both total sulphate and watersoluble sulphate increase with depth. The CEC of the soil is above idme/loog soil, probably due to the presence of montmorillonite. However, the presence of pyrite, which is easily oxidizable to acid sulphate on exposure, makes the soil less suitable for agricultural purposes. INTRODUCTION Acid sulphate soils are those with a ph < 3.5 (:2.5) and with a water soluble sulphate content of >.% within 5 cm of the soil surface 2. The soil contains a high amount of FeS 2 and is normally low in bases. In Peninsular Malaysia alone, acid sulphate soils probably cover an area of about 4,7 ha, found mainly in the coastal region (Fig. ) of Melaka, Kedah, Perlis, Kelantan, Selangor, Trengganu, Johor and Pahang (Law and Seivadurai, 968; Chow, 968; Kanapathy, 966; Paramanatham and Law, 974; Law and Tan, 975; Kanapathy, 976) and are largely grown with padi, coconut, oil palm and rubber. Five soil series namely Telok, Linau, Guar, Sedu and Parit Botak are known to be acid sulphate soils; and are classified at the subgroup level as either sulfic Tropaquept or Typic Sulfaquept (Law and Tan, 975; Gopinathan et al, 977). Acid sulphate soils are derived from marine alluvium. Pvrite in the soils is formed under reducing conditions in the areas intermittently flooded with sea water. Periodic flooding of recent coastal plain soils causes either acidification or deacidification (Breemen, 975). Acidification results in the formation of FeS2 and partial loss of alkalinity in flood water during the wet season, followed by oxidation of pyrite during the dry season. Upon flooding Fe ++ increases at all depths; most strongly in the A horizon. This suggests the occurrence of SO 4 reduction with accompanying FeS 2 prepitation (Breemen and Harmsen, 975). Oxidation of FeS 2 results in the production of acid sulphate and ferric sulphate : FeS 2 oxidation Fe 2 (SO 4 ) 3 Fe 2 (SO 4 ) 3 + 2H 2 O H 2 SO 4 + Fe 2 (SO 4 ) 2 (OH) 2 Present address: Faculty of Education, Universiti Malaya, Kuala Lumpur. 2 ph and sulphate content being measured after drying the soil in the air for 23 weeks. 36

80 SHAMSUDDIN JUSOP AND TUAN OMAR RAJA SENIK Fig. : A map of Peninsular Malaysia shewing the distribution of Quaternary alluvial deposits (shaded area). The dots denote the places where acid sulphate soils are known to occur (adapted from Stauffer, 973). 37

81 The basic ferrisulphate Fe 2 (SO 4 )2(OH}2 so formed can easily be converted to jarosite (K Fe 3 (SO 4 ) 2 (OH) 6 ) in the presence of K^\ Jarosite sometimes dehydrates and oxidizes, resulting in the formation of Fe 2 O 3. These processes are enhanced by a high ph and a low K concentration (Breemen and Harmsen, 975). The object of this paper is to evaluate the agricultural potential of acid sulphate soil in Malaysia by studying mineralogical composition of the soil and physical and chemical properties. MATERIALS AND METHODS ACID SULPHATE SOIL FROM MELAKA The samples were taken at three depths, representing A, (B) and Cg horizons (Fig. 2). This particular Linau Series which is found at Kuala Linggi, Melaka, is derived from Quaternary marine deposits (Stauffer, 973; Sooryanarayana, 976) and largely covered with "gelam" (Melaleuca leucadendron) and or "nipah" (Nipa frutican). Taxonomically, the soil is classified as belonging to the clayey, mixed, isohyperthermic family of the Typic Sulfaquept. The ground water table is below 8 cm depth and yellow jarosite mottles are found in the upper layers in dry season. The dried soil is strongly acid. The mean annual temperature of the area is about 26 C and the total annual rainfall is about 2 25 mm (Dale, 963). H 2 The soil samples were airdried and ground to pass through a 2 mm sieve. Soil ph(h 2 O) was measured at a :2.5 soil. Particlesize analysis was carried out by the pippette method, following the method of Day (965). Exchangeable cations were extracted by NH 4 OAc at ph7 (Chapman, 965); Na and K were determined by a flame photometer and Ca and Mg were determined by atomic absorption spectrophotometer. Exchange capacity was determined by the NH 4 OAc method at ph7 (Chapman, 965). Exchangeable aluminium was extracted by IN KC (Maclean, 965). Time requirement was estimated by the titration method of Dunn (943). Total sulphur was estimated by the method of Jackson (967), sulphatesulphur (SO 4 S) by the method of Hesse (97) and water soluble sulphatesulphur by the method recommended by Chow (968). Organic carbon was determined by the Walkley Black method (Allison, 965). Xray diffraction (XRD) analysis was carried out followig the method of Whittig (965). RESULTS AND DISCUSSION Mineralogical properties Xray diffraction (XRD) investigation of Linau Series (Fig. 3) shows reflection at 6.6A (montmorillinite), 4.A (chlorite), lo.la (illite) 7.2A (kaolinite/chlorite), 3.59A (chlorite), etc (cm) mottle (jarosite) dark brown, humic clay (B) reddish brown, humic clay water table 5 marine clay with peat Fig. 2: A sketch showing the profile of Linau Series from 38 Melaka.

82 SHAMSUDDN JUSOP AND TUAN OMAR RAJA SENIK Cmusc/HUtc quartz halloyslte () kaol (2) nuscoillite (2) NO TREATMENT icoiuite (4) Ksatiirated 6.6 mont. 9.6 *.,K55O C Mgsolvated 78A. Montmorillonite which may be present in the clay does not expand, probably because of the presence of some cementing material which had not been removed by the deferrating agent (Tsuneo, 958). This cementing material was probably destroyed when the Ksaturated clay was heated to 55O C, as a result of which, reflection at 6.6A appears. The 6.6A basal spacing is probably a reflection of montmorillonite. There is no reflection at.8a, thus indicating that halloysite is not present (Whittig, 965). Treated montmorillonite give reflections at about 5A; this shows a high order of crystallinity, which may be attributed to the weathering resistivity of montmorillonite itself. Less or low weathering degree is indicated by the occurrence of chlorite. Mggly. solv. Fig. 3: Xray diffraction pattern of Linau Series saturated with K, K55 C y Mg and Mgglycerol solvated. All these reflections point to the presence of montmorillonite, chlorite, illite and kaolinite. The presence of pyrite was deduced from the presence of the total sulphur and sulphatesulphur. Diffractogram of Mgsaturated and glycerol solvated clay does not manifest reflection at The intensity of reflection at 7.2A, 3.59A and 2.A decreased when heated to a temperature of 55O C. This phenomenon proves the presence of kaolinite as heating causes dehydration and dehydroxylation of kaolinite (Fitzpatrick and Roux, 977). Heating to 55 C does not affect the reflections at 3.4A (illite) and 2.35A and 2.4A (chlorite). There is no clear evidence of the presence of goethite and gibbsite; there are no reflection at 4.86A and 4.8A respectively. The XRD pattern of the upper and lower horizons are basically the same, indicating that there is no real mineralogical difference with depth (Fig. 4); but the increase of cation (Na and Mg) and sulphur with depth points to the corresponding chlo 2.4 musc/illite/quartz / 3.59 kaol Upper horizon (O2O cm) muse illite/halloysite with Lower horizon ( > 5 cm ) H 2 Fig. 4: Xray diffractogram of Ksaturated clays of Linau Series from Melaka. 39

83 increase of montmorillonite and pyrite with depth (Table 2). Mineralogically, the soil may have greater agricultural potential than some upland soils in the country. Montmorillonite increases the exchange capacity of the soil substantially, but the presence of pyrite which is easily oxidizable to acid sulphate, makes the soil less suitable for agriculture. The structure of the soil is poor, but it could be improved provided pyrite is absent. If the pyrite can be destroyed without disturbing the composition of the rest of the component minerals, the soil could be one of the best in the country. However, effective practices for achieving such favourable soil conditions are not available. Draining the soil exposes the pyrites to the air, causing the pyrite to be oxidised to sulphate slowly but steadily over a period of years. Dilution of the acid with large quantities of water and subsequent lime treatment to ph 6.5, is only a short term measure; the remaining pyrite in the soil will eventually be oxidised, rendering the soil unfit for agriculture. Physical and Chemical Properties The texture of the soil becomes heavier with depth; it is sandy clay loam in the A horizon, loam in the (B) horizon and clay loam in the Cg horizon (Table ). ph in H 2 O(:2.5) decreases with depth (Table 2). The decrease in ph is consistent with the increase in sulphate. The acid formed in the A horizon was probably brought down to the lower horizon. Higher ph near the surface may be also due to more water being available in A horizon than in the lower part of the profile. The decrease in ph results in the release of more aluminium into the solution (Table 2). The increase in acidity is subsequently followed by the increase of lime requirement with depth. The presence of a high amount of K in the A horizon is probably due to the presence of jarosite which is yellow in colour (Fig. 2). Potassium decreases downward because jarosite ACID SULPHATE SOIL FROM MELAKA does not form below ground water table. The increase of sodium and magnesium with depth is probably due to the presence of montmorillonite and or the nature of the parent material. The high organic carbon content is attributed to the peaty nature of the Quaternary deposits; pieces of wood are frequently spotted in the Cg horizon (Fig. 2). The CEC of the soil is high, above 6me/ loog soil (Table 2). This is probably due to the presence of montmorillonite. Aluminium and hydrogen dominate the exchange sites at low ph. The presence of the excess amount of Al +++ and H + is undesirable for plant growth. The decrease of ph with depth is small, but the corresponding increase of lime requirement is significantly high. This shows that the soil is highly buffered. High buffering capacity is also shown by the high CEC. The traditional practice of liming the soil under flooded conditions is currently the best form of management for acid sulphate soil. Growing crops which thrive under such conditions is a more logical step than trying to change the soil by artificial means. Under saturated conditions, the most suitable crop is paddy. Flooding acid sulphate soil increases its ph significantly (Ting, 977). The increase in ph is probably brought about by the reduction of hydrous ferric oxide. The ph increase could be due to organic matter which under reduced conditions leads to the accumulation of NH. Ferrous ions are released near the soil surface just above the pyritic substratum (Harmsen and Breemen, 975). Ferrous ions and SO formed in the substratum move upward by diffusion and mass transport. This explains the presence of Fe ++ and SO4S in the upper horizon. Genesis The parent material of Linau Series has probably been subjected to successive lowering and raising of sea level in the Quaternary (Fig. ). Evidence of eustatic rise and fall (due to fluctua TABLE Physical properties of the soil samples Horizon Depth (cm) CSand Particlesize FSand Distribution Silt Clay A (B) Cg >

84 SHAMSUDDIN JUSOP AND TUAN OMAR RAJA SENIK TABLE 2 Chemical properties of the soil samples. Depth (cm) 2 ph (H 2 O) 3.3 Exchangeable Cation (me/loog) Na 2.2 K.33 Ca.5 Mg. Al 2.8 CEC me/loog 7.4 Base Sat(%) 5.5,c 4.23 % Total Sulphur.3 \,SO 4 S.2 %SO4S (H 2 O).4 Lime (ton/ha) > ,, tion of Pleistocene glaciation) of sea level in Peninsular Malaysia are recorded in the numerous drowned mangrove swamps, raised beaches, the prograding stream pattern in the continental shelf (Haile, 97) and punch cores and foraminiferal data (Biswas, 973). Further evidence of sea level rise is seen in the presence of wavecut notches and marine beach deposits around the isolated hill of Bt. Kriang, Kedah, which was once probably an offshore rockly islet (Tjia, 973). It is estimated that the lowest Quaternary sea level was probably about m below and the highest was probably about 4 m above the present one. When sea level was at its maximum, most of the lowland areas (particularly those of Kuala Linggi) where the present acid sulphate soils mostly occur, were submerged (Fig. ). These submerged deposits are dominantly clay and sandy clay with intercalated layers of sand and peat (Stauffer, 973), providing evidence of the existence of both marine and terrestrial deposits. Such a condition was conducive for the accumulation of iron and sulphate, which in reduction resulted in the formation of pyrite when sea level dropped and the deposits were frequently flooded with brackish water. Organic matter provided by 'gelam' and 'nipah' palm helped in the reduction of Fe +++ and SO4. The presence of a high amount of bases, especially those of Na and Mg, under slow leaching (lowlying areas) resulted in the formation of rnontmorillonite. CONCLUSION The study of acid sulphate soil from Kuala Linggi, Melaka, indicates the presence of montmorillonite, kaolinite, illite, chlorite and pyrite. The ph of the soil is less than 3.5 and the CEC is greater than 6me/g soil. Total sulphur, SOJS and water soluble SO4S increase with depth. The presence of montmorillonite and illite followed by high CEC, shows that the soil has great agricultural potential if it was not for the presence of pyrite. Liming is only a short term 4 solution. The subsequent oxidation of pyrite will eventually cause the soil to revert to unfavourable conditions for plant growth. Keeping the soil under anaerobic condition all the time is probably a logical practice under the present circumstances, even though the choice of crops is limited. ACKNOWLEDGEMENTS We gratefully acknowledge the assistance of the Staff of the Department of Soil Science, UPM, in the conduct of the laboratory analysis. We would also like to thank Dr. Othman Yaacob and Encik Khanif Yusop for their helpful suggestions and criticism of the manuscript. REFERENCES ALLISON, L.E. (965): "Determination of Organic Carbon by WalkeyBlack method", Methods of Soil Analysis. (Ed.) Black C A. Am. Soc. Agro BISWAS, B. (973): Quaternary Changes in Sealevel in South China Sea. Bull. geol. Soc. Malays. 6, BREEMEN, N. (975): Acidification and deacidification of coastal plain soils as a result of periodic flooding. Proc. Soil Sci. Soc. Am. 39, BREEMEN, N. and HARMSEN, K. (975): Translocation of iron in acid sulphate soil:i. Soil morphology, and the chemistry and mineralogy of iron in chronosequence of acid sulphate soils. Proc. Soil Sci. Soc. Am. 39, 448. CHAPMAN, H.D. (965): "Determination of cation exchange capacity". Methods of Soil Analysis. (Ed.) Black C.A Am. Soc. Agro. 9, 899. CHOW, W.T. (968): A preliminary study of acid sulphate soils in West Malaysia. Proc 3rd. Malaysian Soil Conf. Malaysian Society of Soil Science, Kuala Lumpur DALE, W.L. (963): Surface temperature of Malaya. J. trop. Geogr. 7, 577. DAY, P.R. (975): Particle fractionation and particlesize analysis. Methods of Soil Analysis. (Ed.) Black C A. Am. Soc. Agro. 9,

85 ACID SULPHATE SOIL FROM MELAKA DUNN, L.E. (943): Lime requirement of soil by means of titration curve. Soil Sci. 56, FITZPATRICK, R.W. and Roux, J.L. (977): Mineralogy and chemistry of a Transvaal Black Clay Toposequence. j'soil Sci. 29, GOPINATHAN, B., PARAMANATHAN, S., NORDIN DAUD, LIM, S.P., KALSI, M.S.andEswARAN, H. (977): Characteristics of some Peninsular Malaysian Soils. CLAMATROPS Conf. Malaysian Society of Soil Science, Kuala Lumpur. HAILE, N.S. (97): Radiocarbon dates of Holocene emergence and submergence in the Tembelau and Bunguran islands, Sunda Shelf, Indonesia. Geol. Soc. Malaysia Bull. 3, HARMSEN, K. and BREEMEN, N. (975): Translocation of iron in acid sulphate soils : II. Production and diffusion of dissolved ferrous iron. Proc. Soil Sci. Soc. Am. 39, HESSE, P.R. (97): A Text Book of Soil Chemical Analysis. London. John Murray Ltd. JACKSON, M.L. (967): Soil Chemical Analysis. New Delhi. Prentice Hall of India, Pte. Ltd. KANAPATHY, K. (966): Acid swamps and problems in their utilization. Paper presented at 2nd Malaysian Soil Conference. Ministry of Agriculture, Malaysia. KANAPATHY, K. (976): Acid Sulphate Soils. Soil and Analylitical Services. Min. Agric. Malays. Bull 6. Kuala Lumpur. LAW, W.M. and SELVADURAI, K. (968): The 968 reconnaissance soil map of Malaya. Proc. 3rd Malaysian Soil Conf. Malaysian Society of Soil Science, Kuala Lumpur LAW, W.M. and TAN, M.H. (975): Range and properties of Peninsular Malaysia soil. Paper presented at the 3rd ASEAN Soil Conf. Ministry of Agriculture, Kuala Lumpur. MACLEAN, E.O. (965): ''Determination of exchangeable aluminium." Methods of Soil Analysis. (Ed.) Black CA. Am. Soc. Agro. 9, PARAMANATHAN, S. and LAW, Y.M. (974): The taxonomy of Peninsular Malaysian soils an approximation. Paper presented at the Symposium: Class. Man. Soils in Malaysia, Malaysian Society of Soil Science, Kuala Lumpur. SOORYANARAYANA, V. (976): A preliminary study of the soils of Melaka. Geographica. : STAUFFER, P.H. (973): "Cenozoic." Geology of the Malay Peninsula. (Ed.) Gobbett DJ. and Hutchison C.S. New York. Wiley & Sons, pp: TING, C.C. (977): Some physicochemical changes on flooding acid sulphate soils. Paper presented at the Annual Fund. Res. Div. Conf. MARDI, Serdang. TJIA, H.D. (973): "Geomorphology". Geology of the Malay Peninsula. (Ed.) Gobbett DJ. and Hutchison C.S. New York. Wiley & Sons, pp: 324. TSUNEO, T. (958): Identification of clay mineral from acid soil. J. Soil Sci. 9(5): 447. WHITTIG, L.D. (965): Xray diffraction technique for mineral identification and mineralogical composition. Methods of Soil Analysis. (Ed.) Black CA. Am. Soc. Agro. 9: (Received 27 June 978) 42

86 Pertanika, (2), 4344 (978) SHORT COMMUNICATION Kesan Nukleotid dan Nukleosid terhadap Aktiviti Fosfodiesterase Mycobacterium smegmatis INTRODUCTION The inhibitory effect of A TP on phosphodiesterase activity plays an important regulatory role in animal cells (Cheung, 966), but this condition has not been demonstrated in microorganisms. This communication shows the inhibitory effect of ATP and other purine and pyrimidine trinucleotides on phosphodiesterase activity in Mycobacterium smegmatis. Although trinucleotides inhibit phosphodiesterase activity, mononucleotides or adenosine stimulate enzyme activity. The results from this study suggest that the intracellular concentration of cyclic AMP is influenced by nucleotides, nucleosides and deoxynucleotides. PENDAHULUAN Kesan penghalangan A TP pada aktiviti fosfodiesterase memainkan peranan pengawalan yang penting dalam selsel haiwan (Cheung, 966), tetapi keadaan ini belum dibuktikan dalam sel mikroorganisma. Kertaskerja ini menunjukkan kesan penghalangan ATP dan Iainlain purin dan pirimidin trinukleotid terhadap aktiviti fosfodiesterase dari bakterium Mycobacterium smegmatis. Walau bagaimanapun trinukleotid menghalang aktiviti fosfodiesterase, kesan stimulasi aktiviti fosfodiesterase boleh didapati dengan mononukleotid atau adenosin. Keputusan dari kajian ini mencadangkan bahawa kepekatan intrasel siklik AMP adalah dipengaruhi oleh nukleotid, nukleosid, dan deoksinukleotid. BAHAN DAN CARA Mycobacterium smegmatis (NCTC 523) dikalcarkan pada suhu 37 C dengan 25 ml larutan medium sintetik dalam ml kelalang konis (Lee, 977). Kalcar bakteria yang ditumbuhkan selama 4 hari dipengemparkan pada 8, g selama 3 min untuk memisahkan larutan medium sintetik dari bakteria. Proses ini diulang dua kali dengan air suling. Kemudian bakteriabakteria diampaikan dengan larutan tampan 6 mm Tris HC (ph 9.) dan didedahkan kepada ayunan sonik selama 2 min pada C untuk memecahkan selsel. Bakteria yang dipecahkan dipengemparkan pada 39 g selama 4 min pada 4 C dan supernatan yang dihasilkan digunakan sebagai punca enzim. Supernatan ini tidak mengandungi enzim adenylate cyclase. Campuran untuk mengesan aktiviti fosfodiesterase mengandungi 2.5 jumol larutan tampan TrisHCl (ph 9.),.5 ^mo\ siklik AMP, PH] siklik AMP (45 cpm), nukleotid atau nukleosid (berbagaibagai kepekatan) dan enzim (4.2 g protein) dalam larutan isipadu 4 //.I, dibiarkan bertindakbalas selama 5 min pada 37 C. Dalam keadaan ini kurang 9% 43 substrat digunakan, penghasilan 5'AMP adalah linear dengan perjalanan masa serta kepekatan enzim, dan aktiviti 5'mononukleotidase tidak boleh dikesan. Tindakbalas ini diberhentikan dengan 5 p} asid trikloroasitik dan tiubtiub uji dipengemparkan pada g selama min untuk memendakkan protein denaturasi. Kemudian 35 pi supernatan digunakan untuk kromatografi kertas (Whatman no. ) dan kertas ini dikembangkan dengan menggunakan sistem solven isopropanol/amonia/air (7::2, isipadu/isipadu). Titiktitik 5'AMP yang jelas diasingkan dari siklik AMP dikesankan dengan lampu ultraungu (254 nm) dan radioaktivitinya diuji dengan penghitung kerlipan. "Recovery" untuk nukleotid yang didapati dari proses kromatografi kertas ialah 85%. Campuran untuk menguji kerlipankerlipan radioaktif mengandungi 2,5difeniloxazole, 4g;,4bis [2(5feniloxazolyl)] benzen.5g; dan toluen dalam larutan isipadu litre. [ 3 H] siklik AMP (6.5 Ci/mmol) yang didapati dari Radiochemical Centre (Amersham, U.K.) ditulinkan dengan menggunakan resin penukaran ion (BioRad AG, X2, 24 mesh, Cl") sebelum digunakan dalam percubaan. Semua nukleotid, nukleosid atau terbitannya didapati dari Sigma Chem. Co. (U.S.A.).

87 C. H. LEE KEPUTUSAN DAN PERBINCANGAN Dengan kehadiran kelebihan 2.5 mm Mg 2 % penghalangan yang penuh terhadap aktiviti fosfodiesterase didapati dengan kepekatan 5 mm ATP. Keputusan ini menunjukkan bahawa ATP adalah penghalang enzim Mycobacteriunt smegmatis. Penghalangan yang penuh oleh ADP hanya sebanyak 87% dalam keadaan kepekatan nukleotid yang tertinggi (Geraf ). reduksi ribonukleotid kepada terbitanterbitan deoksi yang berkenaan (Reichard, Baldesten dan Rutberg, 96). Dalam kajian ini deoksinukleotid pula menghalang aktiviti fosfodiesterase dari M. smegmatis. Walau bagaimanapun penyelidikpenyelidek lain melaporkan bahawa berbagaibagai nukleotid trifosfat menghalang aktiviti fosfodiesterase, keputusan mereka kurang jelas kerana dalam percubaan mereka, kekurangan kepekatan Mg 2 ' sebenarnya menyebabkan penghalangan pada enzim fosfodiesterase. nucleotide (mm) Geraf. Kesan ATP dan ADP pada aktiviti fosfodiesterase. Pada tiaptiap kepekatan nukleotid, kelebihan 2.5 mm Mg l (kecuali yang ditunjukkan) digunakan., ATP;, ATP (dengan tidak kehadiran Mg++);, ADP. Kesan nukleotid dan nukleosid yang lain dikemukakan dalam Jadual. Enzimenzim dari punca mikroba diketahui bahawa ianya memangkin JADUAL Kesan nukleotidnukleotid dan Iainlain sebatian yang berkaitan pada aktiviti fosfodiesterase. Nilai aktiviti enzim dengan tidak kehadiran nukleotid digunakan sebagai " o. Pada tiaptiap kepekatan nukleotid (atau Iainlain sebatian) kelebihan 2.5 mm Mg 2 + digunakan. Kepekatan 2.5 mm Mg2 ^digunakan dalam tiubtiub "Control". Peratus "Control" menunjukkan nilai yang didapati dari purata "triplicate readings". Sebatian 5'GTP (.5 mm) 5'CTP (.5 mm) 5'TTP (.5 mm) 2'deoksi 5'ATP (2. 2'deoksi 5'GTP (2. 2'deoksi 5'CTP (2. 2'deoksi 5'TTP (2. 5'AMP (.5 mm) 5'GMP (.5 mm) 5'CMP (.5 mm) S'TMP (.5 mm) Adenosin (.5 mm) mm) mm) mm) mm) Peratus "Control Fosfodiesterase dari punca bakteria mungkin ujud dalam bentuk metalloprotein seperti yang dicadangkan (Okabayashi dan Ide, 97). Bahanbahan kimia yang bergabung dengan logam enzim akan menghalang aktiviti fosfodiesterase. Dalam kajian ini kelebihan Mg 2+ menentukan bahawa penghalangan oleh ATP tidak boleh diterangkan dengan jelas oleh pengikatan ATP pada logam enzim fosfodiesterase. Mononukleotidmononukleotid dan adenosin merangsang aktiviti fosfodiesterase dari M. smegmatis. Perangsangan adenosin pada enzim fosfodiesterase telah pun dilaporkan dalam tisu buah pinggang (Dousa dan Rychlik, 97). Jabatan Biokimia dan Mikrobiologi Fakulti Sains dan Alam Sekitar Universiti Pertanian Malaysia Serdang, Selangor Malaysia. Lee Chaing Hin PENGHARGAAN Saya mengucapkan terima kasih kepada En. H. Mahmood dan Dr. R. Hinz atas pertolongan mereka. RUJUKAN CHEUNG, W. Y. (966): Inhibition of cyclic nucleotide phosphodiesterase by adenosine 5'triphosphate and inorganic pyrophosphate. Biochem. Biophys. Res. Commun. 23: DOUSA, T., RYCHLIK, I. (97): The metabolism of adenosine 3',5'cyclic phosphate. Biochxm. Biophys. Ada 24: 7.' LEE, C. H. (977): Identification of 3\5* adenosine monophosphate in Mycobacterium smegmatis. J. Bacteriol 32: 333. OKABAYASHI, T., IDE, M. (97): Effect of dipicolinic acid on bacterial cyclic 3',5'nucleotide phosphodiesterase. Biochim. Biophys. Ada 22: REICHARD, P., BALDESTEN, A., RUTBERG, L. (96): Formation of deoxycytidine phosphates from cytidine phosphates in extracts from Escherichia coll J. Biol Chem. 236: 557. (Received 27 October, 978)

88 J^unnouncemen/s INTERNATIONAL CONFERENCE ON AGRICULTURAL ENGINEERING IN NATIONAL DEVELOPMENT, (held in conjunction with the convocation of Universiti Pertanian Malaysia's first batch of Agricultural Engineers). DATE: to 4 SEPTEMBER 979 PLACE: UNIVERSITI PERTANIAN MALAYSIA SERDANG, SELANGOR, MALAYSIA. Papers are invited on the following topics: () The Agricultural Engineering Profession (2) Research and Development in Soil and Water, Mechanisation, Structures and Environment, Agricultural Product Processing and Agricultural Waste Treatment (3) Energy for Agriculture with emphasis on economical use, development and prospects of both conventional and nonconventional sources (4) National Policies The role of Agricultural Engineers in formulating National Policies and Plans for development programmes (5) Implication of Agricultural Engineering in National Development. Summaries of papers should reach the Administrative Secretary before 28 February 978. For further details please write to : The Administrative Secretary, Organising Committee of the International Conference, Faculty of Agricultural Engineering, Universiti Pertanian Malaysia, Serdang, Selangor, MALAYSIA. SYMPOSIUM: LEGUMES IN THE TROPICS (organised by the Faculty of Agriculture, Universiti Pertanian Malaysia) DATE: 3 to 7 NOVEMBER 979 PLACE: UNIVERSITI PERTANIAN MALAYSIA SERDANG, SELANGOR, MALAYSIA. For further information please write to : The Chairman, Organising Committee, Symposium: Legumes in The Tropics, Faculty of Agriculture, Universiti Pertanian Malaysia, Serdang, Selangor, MALAYSIA. 45

89 PUBLICATIONS (UPM PRESS) SEED TECHNOLOGY IN THE TROPICS (35 pages) edited by H.F. Chin, I.C. Enoch and R.M. Raja Harun, Agronomy Department, Faculty of Agriculture, Universiti Pertanian Malaysia. Date published: March 977 'Seed Technology in the Tropics' is composed of papers presented at the first National Seed Symposium 976 in Malaysia. Forty papers by seed technologists from six countries in the tropical and subtropical regions cover the various aspects of seed technology. Discussions at the symposium. The book gives a comprehensive insight and understanding of seed technology in the tropics which undoubtedly differs in various aspects from those of the temperature regions. A useful book for students and seedsmen. Contents cover Breeding; Pests and Protection; Physiology and Storage; Production and Marketing; Testing and Certification; Role of Malaysian Organization in Seed Technology. Price: Hardcover US $. Softcover US$ 7. FOOD AND AGRICULTURE MALAYSIA 2 (53 pages) edited by H.F. Chin, I.C. Enoch and Wan Mohamad Othman, Agronomy Department, Faculty of Agriculture, Universiti Pertanian Malaysia. Date published: May 978 'Food and Agriculture Malaysia 2, is composed of papers presented at the Food and Agriculture Malaysia 2 conference 977 in Malaysia. Over 4 papers presented by specialists in various disciplines such as agronomy, soil science, food technology, economics, nutrition and others from over different countries. Discussions at the symposium. Gives a comprehensive insight into the present day agriculture in Malaysia and future prospects. A useful book for students, planners, agriculturists and food technologists. Contents cover Agriculture, Population and Policies; Climate and Soils; Animal Production; Crop Production; Losses in Food Production; Food Utilization and Consumption; Agricultural Development. Price : Local Overseas Hardcover M$3 US$5 Softcover M$2 US$ A NEW ERA IN MALAYSIAN FORESTRY (323 pages) edited by C.B. Sastry, P.B.L. Srivastava and Haji Abdul Manap Ahmad, Faculty of Forestry, Universiti Pertanian Malaysia. Date published: June 977 'A New Era in Malaysian Forestry, is the proceedings of a Seminar held in March 976. It includes articles by experts and experienced foresters, research scientists and others on different aspects of forestry education, forest science and management, and forest products. Each article is followed by a discussion. The book deals w r ith recent progress, current problems and future trends in forestry education and management of forest resource. It is of interest to students, foresters, conservationists and all those interested in the successful management of tropical forest ecosystems. Articles include: Malaysian Forestry up to the year 2; The Economic Viability of the Malaysian Timber Industry An Overview; The Degree and Diploma Forestry Program at Universiti Pertanian Malaysia; A Contribution of the IBP Research Project to the Understanding of Malaysian Forest Ecology; Preliminary Assessment of Comparative Economics of Plantations and Natural Forests in Malaysia; Designing of Timber Houses in Malaysia; Laboratory Assessment of Pulping Potential of Malaysian Tropical Harwoods; Degradation of Wood by Living Organisms. Price : Local M $5. 46 Overseas US$.

90 Pertanika (2), (978) Author Index for Volume A Abdul Ghani Ibrahim., 55 Awaluddin Hj. Talib., 7 Mohamad Idris Haji Zainal Abidin., 7 Mohamed Ismail Abdul Karim., 4 Mohd Noor Ismail., 7 Mohd Yusof Hussein., 36 Mohd Rashdan bin Haji Baba., B Baharin Kassim., 97 Beilharz, E.G., 97 Ng Eng Gim., 7 N Cheok, K.T., 82 C Othman Yaacob., 7 O D Daud Khamis., 26 Rosli Mohamad., 66 K Khor Geok Lin., 7 Kiew, Ruth., 2 Kuan, K.K., 62 Shamsuddin Jusop., 36 Srivastava, P.B.L., 26 Law, A.T., 5 Lee, C.H., 43 Lim Eng Siong., 59 Lim, G.S., 2 Tan, S.H., 3 Tan, S.T., 3 Tan, S.G., 22, 86 Tuan Omar Raja Senik., 36 M Mak, T.K., 62 Mohammed ArifT Hussein., 5 Yusof Ibrahim., 66 Yap, T.C., 3, 82 ISSN

91 Pertanika (2), 4852(978) Subject Index for Volume ATP Kesan penghalangannya terhadap aktiviti f osf odiesterase 43 Acetocarmine stain to stop growth of pollen tubes 59 Achrotichium aureum 27 Acid sulphate soil mineralogy and related properties of, from Melaka, Malaysia 36 Adenilat kinase (AK) see Genetic markers, biochemical Adenosin see Adenosine Adenosin deaminase (ADA) see Genetic markers, biochemical Adenosine meransang aktiviti fosfodiesterase 44 Agar culture medium for pollen germination 59 Albumin see Genetic markers, biochemical Alocasia aft. macrorrhiza (L.) Schott ground flora of Gunung Mulu National Park, Sarawak 3 Aristichthys nobilis see Big head 5 Asparagus bean see Long bean Artocarpus heterophyllus (jackfruit) 66 Averrhoa carambola (starfruit) 66 Bacterial fermentation to neutralize excess alkalinity of potato waste 4 optimun conditions for 4 Bean fly pest of leguminous plants in Malaysia 36 Begonia ground flora of Gunung Mulu National Park, Sarawak 4 Big head cultured under the polyculture system 5 Birth weight differences among Landrace, Duroc and their crossbreds 62 Blood groups datadata bagi pendudukpenduduk Malaysia, Brunei & Singapore 86 Blowfly leg paralysis induced by Risella 7 oil 26 Brabender instruments used for rheological evaluation of dough in breadmaking 8 Bread characteristics of loaf volume organoleptic 3 shelf life of 4 Bread, composite 8 48 Bread, control (standard) 8 Bruguiera cylindrica 27 Bruguiera gymnorrhiza 28 Bruguiera parviflora 28 Bruguiera sexangula 28 Bruguiera spp. 27 Carica papaya (papaya) 66 Calanthe ground flora of tropical lowland rain forest, Sarawak 6 Carbofuran effect on plant height 38 to control beanfly by soil application 36 Cassava replacing wheat flour for breadmaking 7 Cation exchange capacity (CEC) of acid sulphate soil 4 Chinta (sweet corn) see Maize Coconut supply and demand 6 Corn see Maize Correlation coefficient genotypic and phenotypic of stalk crushing strength and grain yield of maize 9 Costus speciosus (Koenig) ground flora of Gunung Mulu National Park, Sarawak 3 Ctenopharyngodon idella Valenciennes see Grass carp Curculigo latifolia Dryand ground flora of Gunung Mulu National Park, Sarawak 3 Cyrtandra ground flora of Gunung Mulu National Park, Sarawak 3 Cyrtandra oblongifolia Bl ground flora of Gunung Mulu National Park, Sarawak 5 Cyrtandra pendulifera ground flora of Gunung Mulu National Park, Sarawak 7 Cyrtandra radicifolia C. B. Cl ground flora of Gunung Mulu National Park, Sarawak 7 Dacus dorsalis see Fruit fly Derris uliginosa 27 Digestion of the Letnna spp. (duckweed), Guinea grass with Napier grass as food in grass carp 52

92 Donax canniformis Forst. ground flora of Gunung Mulu National Park, Sarawak 3 Driessenia ground flora of Gunung Mulu National Park, Sarawak 3 Durian clones 7 observation on growth and early production 7 Durio zibethinus Mur see Durian Duroc see Pigs Elatostema acuminata (Poir.) Brongn. ground flora of Gunung Mulu National Park, Sarawak 5 Environment, Genotype role in maize 6 Epithema involucratum Roxb. ground flora of Gunung Mulu National Park, Sarawak 3 Euthemis ground flora of Gunung Mulu National Park, Sarawak 3 Exoecaria agollochia 28 Favism penyakit dari kekurangan aktiviti enzim eritrosit gluko6fosfat dehidrogenase 32 Fertiliser basic requirements and standard practice for durian cultivation 73, 75 Flora ground flora of tropical rain forest 7 Flour, composite utilizing local carbohydrate rich resources 7 Flowers flower and fruit drop of durian clones in Universiti Pertanian Malaysia 8 flowering and senescence during fruiting period of long bean 83 flowering pattern of various durian clones 7 Foliage, iridescent and variegated of ground flora at Gunung Mulu National Park, Sarawak 3 Forest floristic components of the ground flora of a tropical lowland rain forest 2 see also Mangrove Fosfatase asid (air liur) see Genetic markers, biochemical Fosfodiesterase see Phosphodiesterase Fosfoglukonat dehidrogenase (6PGD) see Genetic markers, biochemical Fosfoglukomutase I (PGM) see Genetic markers, biochemical French bean effect of bean fly 36 Fruit flower and fruit drop of durian clones in Universiti Pertanian Orchard 8 49 Fruit fly pupal distribution among fruit trees 66 pupation depth among fruit trees 67 Genesis of acid sulphate soil in Peninsular Malaysia 4 Genetic markers dalam orang asli semenanjung Malaysia, Sabah, Sarawak and Brunei 86 Genetic markers, biochemical dari eritrosit, data bagi penduduk Malaysia 27,9 mengikut air liur, data bagi penduduk Malaysia 26, 9 mengikut jenis serum, data bagi penduduk Malaysia 23, 9 mengikut varian, data bagi penduduk Malaysia 3, 93 Gini Concentration Ratio for measuring equity in Peninsular Malaysia 5 Glukos6fosfat dehidrogenase (G6PD) see Genetic markers, biochemical Glysine max see Soya bean Gram positive cocci microorganisms found in fermented waste see microorganism Gram positive rods microorganisms found in fermented waste see microorganism Grass carp digestibility of Napier grass in 5 Growth pattern of durian (Durio zibethinus Murr) trees, in comparison among different clones 7 Hanguana malayana Merr. ground flora of Gunung Mulu National Park, Sarawak 4, 6 Haptoglobin see Genetic markers, biochemical Hemoglobin (Hb) see Genetic markers, biochemical Heterosis yield performance in maize 3 Heritiera littoralis 28 Hexatheca fulva C. B. Cl. ground flora of Gunung Mulu National Park, Sarawak 3, 4 Homalomena humilis ground flora of Gunung Mulu National Park, Sarawak 4, 6 Hoseanthus lobbii ground flora of Gunung Mulu National Park, Sarawak 3 Hypophthalmichthys molitrix see Silver carp 5 Illite in acid sulphate soil 39

93 Iguanura melinauensis Kiew ground flora of Gunung Mulu National Park, Sarawak 8 Income estimates for rubber, coconut and pineapple small holders in Peninsular Malaysia 7 Insecticides Risella 7 oil used in insecticide formulations 2 carbofuran see Carbofuran Irish potato see Potato Kerangas forest see Kumpulan darah see Forest Blood groups Landrace see Pigs Lasianthus stipularis ground flora of Gunung Mulu National Park, Sarawak 7 Lemna spp. see Duck weed Lepidagathis longifolia ground flora of Gunung Mulu National Park, Sarawak 7 Linau series soil see Acid sulphate soil Long bean evaluation of using early and late produced seeds in two soils 82 commercial varieties grown in Malaysia 82 Lucilia sericata Meig see Blowfly Macrobrachium rosenbergii see Udang galah Maize hybrid resistant for stalk strength 7 new composite sweet corn 4 Recurrent Selection Methods 3, 8 sweet corn 3 Malaxis ground flora of Gunung Mulu National Park, Sarawak 8 Manihot esculenta see Cassava Mangrove forests composition of seedling crop 28 current silvicultural practices in Matang Mangrove Reserve 26 mortality of seedlings 33 natural regeneration 28 2 months after felling months after felling 34 progress of natural regeneration 26 Matang Mangrove Reserve see Mangrove forests Mapania cuspidata ground flora of Gunung Mulu National Park, Sarawak 6 Mice bodyweight at three and nine weeks, 2 coefficient of inbreeding 98 effects of selection for highlitter weight on reproductive performance 97 5 littersize at birth 99 litterweight at three and nine weeks mortality rate among selected lines 99, percentage of successful mating 98 selection and inbreeding 98 Microflora source of bioprotein 52 role in the digestion of grass in fish 53 Microorganism gram positive rods and cocci in acid production in fermented potato waste 42 Mononucleotides merangsang aktiviti fosfodiesterase 43 Mononukleotid see Mononucleotides Montmorillonite in acid sulphate soil suitable for agriculture 4 Mortality of blowfly, the effect of Risella 7 oil 2 Mung bean effect of carbofuran on 36 Musa campestris ground flora of Gunung Mulu National Park, Sarawak 3, 4 Mycobacterium smegmatis pengawalan metabolisma siklik MP dalam sel 43 Myrmeconauclea strigosa ground flora of Gunung Mulu National Park, Sarawak 3, 4 Napier grass digestibility in grass carp 5 Natural rubber supply and demand 6 Neckia serrata ground flora of Gunung Mulu National Park, Sarawak 4, 7 Ophiomyia phaseoli Tyron see Bean fly 36 Pennisetum purpureum see Napier grass Perspex block in the study of pollen growth 59 Peptidase B (PEP B) see Genetic markers, biochemical Phaseolus vulgaris see French bean Photosynthates effect of drought; and affecting stalk crushing strength and yield 2 Phosphodiesterase kesan penghalangan nukleotid dan nukleosid terhadap aktivitinya 4 Phyllagathia elliptica ground flora of Gunung Mulu National Park, Sarawak 4, 6 Pigs estrus and estrus cycles of Landrace and Duroc gilts 63

94 exotic breeds in Malaysia Landrace, Duroc, Yorkshire Hampshire & Chesterwhite 62 management conditions of gilts for crossbreeding 63 observations on Landrace, Duroc and their crossbreds 62 Pineapple semipermanent crop in Malaysia 6 Piptospatha elongata ground flora of Gunung Mulu National Park, Sarawak 3 Pirimidin trinukleotid see Pyrimidine trinucleotide Plant height effect of beanfly 37, 38 Pod length analyses of variance in long bean varieties 83 effect of environmental changes 83 Pollen germination 59 Polyathia flagellaris ground flora of Gunung Mulu National Park, Sarawak 4 Potato lye peeled white or Irish potato waste as feedstuff 4 Price elasticity of natural rubber, coconut and pineapple 6 Psidium guajava (guava) 66 Psophocarpus tetragonolobus see Winged bean 8 Purin see Purine Purine kesan penghalangannya terhadap aktiviti fosfodiesterase 43 Pyrimidine trinucleotide kesan penghalangannya terhadap aktiviti fosfodiesterase 43 Pyrite in acid sulphate soil not suitable for agriculture 4 Rainfall at Serdang 7 Randomized Complete Block Design Pattern for population allocation of maize 4 in evaluation of long bean lines 82 Recurrent Selection Methods for further yield improvement in maize 3 to improve stalk quality in maize 8 Rhizophora apiculata 28, 33 Rhizophora mucronata 28 Rhizophora spp. 27 Si Progeny Tests to determine stalk strength and grain yield in maize 7 Selaginella ground flora of Gunung Mulu National Park, Sarawak 3, 4 5 Silver carp cultured under the poly culture system 5 Smallholders of rubber, coconut, pineapple in Johore, Malaysia 5 Soils Acid sulphate soil for Melaka, Malaysia mineral and related properties 36 mechanical and chemical composition of soil in durian orchard of Universiti Pertanian Malaysia 7 Solarium tuherosum see Potato Sonerila puchella ground flora of Gunung Mulu National Park, Sarawak 7 Soya bean effect of carbofuran 36 Stalks relationship of stalk strength and grain yield in maize 7 Stachyphyrnium ground flora of Gunung Mulu National Park, Sarawak 7 Sucrose for pollen germination 59 Sulfic tropaquept see acid sulphate soil Superoksid dismutase B (SOD B) see Genetic markers, biochemical Sweet corn see Maize Tandatanda genetik biokimia see Genetic markers, biochemical Tax burden on rubber, coconut and pineapple small holders in Johore 5 Tax incidence definition and incidence assumption in developing countries 6 Taxes education tax on small holders in Peninsular Malaysia 8 export duty on rubber, coconut and pineapple 9 import taxes on inputs used in rubber, coconut and pineapple production 7 land tax paid by small holders in Malaysia 8 Transferrin see Genetic markers, biochemical Udang galah cultured under the polyculture system 5 Vicia faba L. pollen from fresh flowers of 59 Vigna sisquipedalis see Long bean Vigna radiata see Mung bean Vigna sisquipedalis Fruw see Long bean

95 Water Winged been effect on yield and stalk crushing strength in as protein suplement of cassavewheat bread maize 9 (composite bread) 8 Weaning weight diffraction d.fferences among Landrace, Duroc and their crossbreds 63 > ^ analv8fe Qn Hnau series ^ 3g Weeding Yield effect of weeding with implements driven by relationship of stalk strength and grain yield heavy tractors on durian trees 8 in maize 7 White or Irish potato see Potato Yard long bean see Long bean 52

96 Pertanika (2), (978) ACKNOWLEDGEMENT The Editorial Board acknowledges in the preparation of Volume. Dr. A. Aziz b. S. A. Kadir Dr. Abdul Latif b. Ibrahim Dr. Ahmad Zaharuddin Idrus En. N.T. Arasu Prof. Amudi Pasaribu Dr. Baharin Kassim Dr. A. Balasubramaniam En. L.C. Cheah Dr. C. Devandra Prof. I.C. Enoch Dr. CO. Fong Prof. J.I. Furtado Dr. Harun Deruah Dr. Khalid Hassan Prof. Madya Kardinal Kusnaeny the help of the following reviewers Puan L.L. Lim Dr. S.C. Lim Prof. Mohd. Zain Hj. Karim Dr. T.K. Mukherjee Dr. F.S.P. Ng Dr. K.K. Ong Dr. E. Pusparajah Dr. K.G. Singh Prof. E. Soepadmo Dr. LK. Sudderudin Dr. B.T. Tan En. P.O. Thomas En. Y.P. Tho Dr. Bobby Tee Dr. T.C. Yap Dr. E.S. Lim 53

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