REINWARDTIA A JOURNAL ON TAXONOMIC BOTANY, PLANT SOCIOLOGY AND ECOLOGY Vol. 14(1): 1 - 2 4 8 , December 23, 2014 Chief Editor Kartini Kramadibrata (Mycologist, Herbarium Bogoriense, Indonesia) Editors Dedy Darnaedi (Taxonomist, Herbarium Bogoriense, Indonesia) Tukirin Partomihardjo (Ecologist, Herbarium Bogoriense, Indonesia) Joeni Setijo Rahajoe (Ecologist, Herbarium Bogoriense, Indonesia) Marlina Ardiyani (Taxonomist, Herbarium Bogoriense, Indonesia) Topik Hidayat (Taxonomist, Indonesia University of Education, Indonesia) Eizi Suzuki (Ecologist, Kagoshima University, Japan) Jun Wen (Taxonomist, Smithsonian Natural History Museum, USA) Managing Editor Himmah Rustiami (Taxonomist, Herbarium Bogoriense, Indonesia) Lulut Dwi Sulistyaningsih (Taxonomist, Herbarium Bogoriense, Indonesia) Secretary Endang Tri Utami Layout Editor Deden Sumirat Hidayat Medi Sutiyatno Illustrators Subari Wahyudi Santoso Anne Kusumawaty Correspondence on editorial matters and subscriptions for Reinwardtia should be addressed to: HERBARIUM BOGORIENSE, BOTANY DIVISION, RESEARCH CENTER FOR BIOLOGY- INDONESIAN INSTITUTE OF SCIENCES CIBINONG SCIENCE CENTER, JLN. RAYA JAKARTA - BOGOR KM 46, CIBINONG 16911, P.O. Box 25 Cibinong INDONESIA PHONE (+62) 21 8765066; Fax (+62) 21 8765062 E-MAIL: reinwardtia@mail.lipi.go.id 1 2 3 4 1 3 4 4 Cover images: 1. Begonia holosericeoides (female flower and habit) (Begoniaceae; Ardi et al.); 2. Abaxial cuticles of Alseodaphne rhododendropsis (Lauraceae; Nishida & van der Werff); 3. Dipo- dium puspitae, Dipodium purpureum (Orchidaceae; O'Byrne); 4. Agalmyla exannulata, Cyrtandra coccinea var. celebica, Codonoboea kjellbergii (Gesneriaceae; Kartonegoro & Potter). The Editors would like to thanks all reviewers of volume 14(1): Abdulrokhman Kartonegoro - Herbarium Bogoriense, Bogor, Indonesia Altafhusain B. Nadaf - University of Pune, Pune, India Amy Y. Rossman - Systematic Mycology & Microbiology Laboratory USDA-ARS, Beltsville, USA Andre Schuiteman - Royal Botanic Gardens, Kew, UK Ary P. Keim - Herbarium Bogoriense, Bogor, Indonesia Barry Conn - Royal Botanic Gardens National Herbarium of New South Wales, Sydney, Australia Dato' Abdul Latiff Mohamad - Universiti Kebangsaan Malaysia, Bangi, Selangor, Malaysia Daniel Potter - Department of Plant Sciences, University of California, Davis, California, USA Deby Arifiani - Herbarium Bogoriense, Bogor, Indonesia Ferry J. W. Slik - University of Brunei Darussalam, Brunei Henti H. Rachmat - Conservation and Rehabilitation Research and Development Center, Bogor, Indonesia Ian M. Turner - Royal Botanic Gardens, Kew, UK Iskandar Z. Siregar - Bogor Agricultural University, Bogor, Indonesia Jay H. Bernstein - Kingsborough Community College, Brooklyn, New York, USA Jens G. Rohwer - University of Hamburg, Hamburg, Germany Joan Pereira - SAN Herbarium, Sabah Forestry Department, Sabah, Malaysia Kuswata Kartawinata - Herbarium Bogoriense, Bogor, Indonesia Lars H. Schmidt - University of Copenhagen, Copenhagen, Denmark Mark Hughes - Royal Botanic Gardens, Edinburgh, UK Masahiro Kato - Kyoto University, Kyoto, Japan Nuril Hidayati - Herbarium Bogoriense, Bogor, Indonesia Ong Poh Teck - Forest Research Institute Malaysia, Kepong, Malaysia Peter C. van Welzen - National Herbarium Netherlands, Leiden University Branch, Leiden, Netherlands Reuben Nilus - Sabah Forestry Department, Sabah, Malaysia Rugayah - Herbarium Bogoriense, Bogor, Indonesia Ruth Kiew - Forest Research Institute of Malaysia, Kepong, Malaysia Uwe Braun - Institut fur Biologie Bereich Geobotanik und Botanischer Garten, Halle (Saale), Germany Yasuaki Sato - Osaka-Sangyo University, Osaka, Japan REINWARDTIA Vol 14, No 1, pp: 237 − 247 237 grow in rainforests worldwide (Dransfield, 1978). In Australia, approximately 60% of palm species occur in rainforest habitats (Dowe, 2010). Arenga australasica (H. Wendl. & Drude) S. T. Blake ex INTRODUCTION Palms are an important component of many tropi- cal rainforests. Approximately 75% of palm species A PHYSIOLOGICAL APPROACH TO CONSERVATION OF FOUR PALM SPECIES: ARENGA AUSTRALASICA, CALAMUS AUSTRALIS, HYDRI- ASTELE WENDLANDIANA AND LICUALA RAMSAYI Received January 13, 2013; accepted October 23, 2014 DIAN LATIFAH Center for Plant Conservation-Botanic Gardens, Indonesian Institute of Sciences (LIPI) Jln. Ir. H.Juanda no. 13, Bogor, Indonesia 16122. E-mail: latifah2311@yahoo.com ROBERT A. CONGDON College of Marine & Environmental Sciences, James Cook University, James Cook Drv, Townsville, Queensland, Australia 4811. E-mail: robert.congdon@jcu.edu.au JOSEPH A. HOLTUM College of Marine & Environmental Sciences, James Cook University, James Cook Drv, Townsville, Queensland, Australia 4811. E-mail: joseph.holtum@jcu.edu.au ABSTRACT LATIFAH, D., CONGDON, R. A. & HOLTUM, J. A. 2014. Physiological approach of four palm species conservation: Arenga australasica, Calamus australis, Hydriastele wendlandiana and Licuala ramsayi. Reinwardtia 14(1): 237 – 247. ― Palms (Arecaceae) are an important component of many tropical rainforests. Many have also been cultivated widely for agricultural commodities with high economic value. They are also important components in rehabilitation of disturbed or marginal lands. Knowledge and application of germination strategies are essential in the cultivation of palms. Many species have seeds that do not germinate readily, even when light conditions are favourable. This research determined the effects of seed coats, light and temperature on germination of Arenga australasica (H. Wendl. & Drude) S. T. Blake ex H. E. Moore, Calamus australis Mart., Hydriastele wendlandiana (F. Muell.) H. Wendl. & Drude and Licuala ramsayi var. tuckeri Barford & Dowe. We examined physical treatments to promote germination or break dormancy, as well as different light and temperature conditions. The results showed that the hard seed coats of the four species slowed imbibition. Scarified seeds germinated best for A. australasica, C. australis and L. ramsayi. The germi- nation of all seeds was inhibited by far red light. The red light requirement suggests that these species prefer to colonise open areas. This implies that dispersal agents, canopy gaps and forest margins may play important roles in promoting regeneration as well as conservation of these palm species. Key words: Arenga australasica, Calamus australis, germination, Hydriastele wendlandiana, Licuala ramsayi, palms. ABSTRAK LATIFAH, D., CONGDON, R. A. & HOLTUM, J. A. 2014. Pendekatan fisiologi pada konservasi empat jenis palem: Arenga australasica, Calamus australis, Hydriastele wendlandiana dan Licuala ramsayi. Reinwardtia 14(1): 237 – 247. ― Palem (Arecaceae) adalah komponen penting di hutan hujan tropis. Banyak jenis yang dibudidaya secara luas untuk komoditi pertanian dengan nilai ekonomi yang tinggi. Selain itu palem juga penting untuk merehabilitasi lahan-lahan marginal atau terganggu. Pengetahuan serta aplikasi strategi perkecambahan sangat penting dalam pembudidayaan palem. Banyak jenis yang mempunyai biji yang tidak langsung berkecambah walaupun dalam kondisi cahaya yang sesuai. Penelitian ini bertujuan untuk mengetahui pengaruh pelindung biji, cahaya dan suhu pada perkecambahan Arenga australasica (H. Wendl. & Drude) S. T. Blake ex H.E. Moore, Calamus australis Mart., Hydriastele wendlandiana (F. Muell.) H. Wendl. & Drude dan Licuala ramsayi var. tuckeri Barford & Dowe. Karakter fisik telah diamati untuk merangsang perkecambahan atau memecah dormansi demikian juga dengan perbedaan cahaya dan kondisi suhu. Hasil penelitian menunjukkan bahwa pelindung biji yang keras menunjukkan imbibisi. Perkecambahan biji terbaik pada A. australasica, C. australis dan L. ramsayi. Perkecambahan kesemua jenis palem tersebut banyak dipengaruhi oleh cahaya merah. Kebutuhan akan cahaya merah menunjukkan bahwa kesemua jenis tersebut lebih menyukai daerah terbuka. Hal ini berimplikasi pada agen penyebar, perbedaan kanopi serta perbatasan hutan yang kemungkinan mempengaruhi daya regenerasi dari jenis palem tersebut. Kata kunci: Arenga australasica, Calamus australis, Hydriastele wendlandiana, Licuala ramsayi, palem, perkecambahan. REINWARDTIA 238 [VOL.14 H. E. Moore, Calamus australis Mart., Hydriastele wendlandiana (F. Muell.) H. Wendl. & Drude and Licuala ramsayi var. tuckeri (F. Muell.) Domin have not been assessed for the IUCN Red List (IUCN, 2014). The conservation status of A. aus- tralasica is vulnerable (EPA, 2007), whereas, the other study species are not currently threatened (Dowe, 2010). Furthermore, A. australasica is endemic to north-eastern Queensland and the adjacent islands (Dowe, 2010); C. australis is endemic to Queensland (Hyland et al., 2003); H. wendlandiana is endemic to Queensland and the Northern Territory (Dowe, 2010; Hyland et al., 2003); and Licuala ramsayi var. tuckeri is restricted to north and north eastern Queensland (Barfod & Dowe, 2005; Dowe, 2010). Therefore, the conserva- tion of these species is an important issue. Studies have shown that canopy gaps play an important role in stimulating recruitment in most plant communities (Bullock, 2000; Fenner & Thompson, 2005; 1977). This may be caused by reduced competition for resources, such as light, nutrients and water. Bullock (2000) differentiated three stages of seedling development that may be influenced by canopy gaps: ‘germination’, ‘emergence’ and ‘establishment’. Detecting early germination stages in the field can be difficult, but ‘emergence’ can be readily seen. Hence, most studies of germination responses to canopy gaps require the use of artificial shading in laboratories or glasshouses to compare to “full light” responses (Bullock, 2000). This study used laboratory trials on seed germination to determine the effects that changed light conditions are likely to have on palm regeneration following natural disturbance. Many palms have seeds that do not germinate readily, even when light conditions are favourable. This seed dormancy is commonly caused by the hard seed coat that restricts the exchange of water and gases (Dessai & Salunkhe, 1997). It is also well known that palm seed dormancy is caused by a combination of immature embryos and the failure of developing embryos to rupture covering structures (Pérez, 2009; Pinheiro, 2001; Pritchard et al., 2004). Dormancy is the suspension of growth by active endogenous inhibition (Jann & Amen, 1977), or “a state in which a viable seed will not germinate when placed in conditions normally considered to be adequate for germination, that is, when provided moisture and oxygen” (Tran & Cavanagh, 1984), and with suitable temperature. However, some conflicts in defining seed dormancy have arisen. Fenner & Thompson (2005) reviewed the conflict- ing views of ecologists and physiologists on seed dormancy. They suggested integrating the conflict- ing views by following Vleeshouwers et al. (1995) who have argued that “(1) dormancy should not be identified with the absence of germination, and (2) dormancy is a characteristic of the seed (and not the environment) and defines the conditions necessary for germination”. Three types of dormancy are recognised: (1) in- nate, (2) induced and (3) enforced (Harper, 1977). Other researchers, Adkins and Bellairs (2000) di- vide dormancy mechanisms into two types: (1) “coat-imposed dormancy”, and (2) “embryo dor- mancy”. The dormancy of some palm seeds may be classified as ‘hard seed coat-imposed dormancy’. This type of dormancy presents in the seeds of Cyr- tostachys lakka and Roystonea elata (Soedjono & Suskandari, 1996). Dormancy in these two palm species was broken by mechanically scarifying seeds by filing. Embryo dormancy may also present in some palm seeds. Dormancy in Butia spp. appears to be caused by an immature embryo (Sento, 1986). The embryo immaturity, combined with the failure of developing embryos to release a covering structure, i.e. operculum, led to ‘morpho-physiological’ dor- mancy in Pritchardia remota seeds (Pérez, 2009; Pérez et al., 2008). There are two methods of breaking dormancy: either through natural break- down or artificial treatments (Koch & Dixon, 2000). Previous observations on the study species are pre- sented in Table 1. The literature reveals few de- tailed studies of germination in these species. No information could be found on dormancy cha- racteristics and the observations on germination of H. wendlandiana are inconsistent. This research used laboratory trials to determine the effects that changed environmental conditions are likely to have on regeneration of palms follow- ing cyclonic disturbance. These experiments inves- tigated the effects of seed coat, light and tempera- ture on germination, with the following hypotheses: (i) Germination will vary between species under three treatments: (1) no treatment; (2) moist heat at 50ºC; and (3) scarification. This will imply that seed germination is affected by hard seed coats that inhibit water absorption. (ii) Germination will vary between species under three treatments: (1) 20ºC and continuous white light; (2) 30ºC and continuous white light; and (3) alternating light at 30°C and dark at 20°C. Treat- ments were chosen to reflect the effect of conditions under a forest canopy and in canopy gaps or diurnal/diel changes of light and temperature on germination. (iii) Germination will vary between species under five treatments: (1) continuous white light; (2) dark- ness; (3) continuous red light; (4) continuous far red light; and (5) a combination of red and far red light. 2014] 239 LATIFAH et al.: A physiological approach to conservation of four palm species The results will indicate whether germination is af- fected by the quality of light passing through the canopy (far red/shorter red light, 730 nm spectrum, Atwell et al., 1999), in canopy gaps (red light, 660 nm spectrum, Atwell et al., 1999), or the proportion of red to far red light. MATERIAL & METHODS Seeds of A. australasica and H. wendlandiana were obtained from mature specimens in the Townsville Palmetum, C. australis from Tam O’Shanter National Park at Mission Beach, H. wendlandiana from Kurrimine Beach National Park, and L. ramsayi from the Anderson Park Botanic Garden, Townsville. Longitudinal sections of the seeds demonstrate the nature of the seed coats (Fig. 1) and seed dimensions are given in Table 2. The seeds were fully imbibed to activate the seed metabolism for germination (ISTA, 1985). After the treatments, Thiram fungicide (active constituent: 800g/kg Thiram) was applied and all seeds were placed in a 20 or 30ºC controlled-temperature room under the relevant light conditions depending on theexperiment (Table 3). The source of continuous white light was a fluorescent lamp (36W/840 cool white Philips Lifemax). A few containers full of water were placed in various positions in the rooms to maintain humidity. A completely randomized research design was applied to address the hypotheses (Gasperz, 1991). The dormancy breaking experiment used artificial techniques to simulate natural conditions that tend to take a long time. Therefore, for example, seeds were scarified by filing the seed coat deep enough A B C D Table 1. Summary of previous observations on germination of the five palm species examined in this study. (n.a. de- notes information is not available). Fig. 1. Longitudinal sections of seeds of A. A. australasica, B. C. australis, C. H. wendlandiana, D. L. ramsayi showing a = apical end, b = testa/seed coat or sarcotesta in C. australis (Swartz, 1971; Uhl & Dransfield, 1987), c = endosperm, d = embryo, e = fibrous sheath (Essig & Hernandez, 2002), f = endocarp. Scale is in mm. Species Germination type Information on germination A. australasica (H. Wendl. & Drude) S. T. Blake ex H.E. Moore Remote non ligular (Latifah, 2011) 90% of the seeds germinate within 19 days at 30°C; details of germination methods were not explained1 (ISTA, 2006); dormancy occurred in the related species A. microcarpa (Latifah, 2004). C. australis Mart. adjacent ligu- lar in Cala- mus genera (Dowe, 2010) The germination capability of other Calamus spp varies: C. peregrinus showed 91% germination within 12-35 days with sarcotesta removed (Vangkualong, 1984); heat treatment at 40 ºC for 48 hours in C. latifolius increased germination (Mohd et al., 1994); C. mannan and C. caesius germinate within 2-3 weeks after sowing in nursery beds consisting of a sand:soil mix (1:4 or 1:3) and sown seeds were covered by 3 cm of sawdust (Ahmad & Hamzah, 1984); dormancy: n.a. C. moti F.M. Bailey H. wendlandi- ana (F. Muell.) H. Wendl. & Drude n.a The seeds need scarification to germinate after 365 days (Lawie, 2007). Fresh seeds can also germinate within 3-6 months (Cronin, 1989); dormancy: n.a. L. ramsayi var. tuckeri Barford & Dowe remote- ligular in Licuala gen- era Uhl & Dransfield (1987) The fresh seeds of L. ramsayi germinate after 6 months on various rich, well- drained soil types (Cronin, 1989); dormancy: n.a. REINWARDTIA 240 [VOL.14 Table 2. Seed dimensions of study species. L = length (mm), W = width (mm) and D = diameter (mm), for seeds that are round in cross-section. Table 3. Overview of the germination experiments in four palm species including the details of each experiment and treatment. The table also presents the number of replicates and seeds used in each experimental unit; r= number of replicates; s= number of seeds in each replicate; n.a.=seeds were not available. There were insuf- ficient seeds of H. wendlandiana which restricted the number of light treatments used. No seeds of C. moti were available for these experiments. Species N Weight (g) Dimension (mm) Mean Range Mean Range A. australasica 127 0.75±0.02 0.20-1.15 L=12.92±0.31 W=10.33±0.21 L=8.73-17.31 W=7.79-12.79 C. australis 25 0.62±0.03 0.43-0.68 L= 10.05±0.22 D=10.33±0.17 L= 8.73-10.89 D= 9.31-10.86 H. wendlandiana 55 0.07±0.004 0.04-0.08 L=6.63±0.22 D=4.50±0.13 L=5.52-7.71 D=3.73-5.06 L. ramsayi var. tuckeri 50 0.27±0.02 0.13-0.34 D=7.04±0.17 D=5.66-7.58 Experiments A. australasica C. australis H. wendlandiana L. ramsayi Experiment 1: Effect of Dormancy Breaking Treatments on Imbibition (1) No treatment n = 4r x 10s n = 4r x 10s n = 3r x 7s n = 5r x 13s (2) Scarification n = 4r x 10s n = 4r x 10s n = 3r x 7s n = 5r x 13s (3) Moist heat 50°C n = 4r x 10s n = 4r x 10s n = 3r x 7s n = 5r x 13s Experiment 2: Effect of Dormancy Breaking Techniques on Germination (1) No treatment n = 4r x 10s n = 4r x 10s n = 3r x 7s n = 5r x 13s (2) Scarification n = 4r x 10s n = 4r x 10s n = 3r x 7s n = 5r x 13s (3) Moist heat 50°C n = 4r x 10s n = 4r x 10s n = 3r x 7s n = 5r x 13s Experiment 3: Diel Fluctuation of Light and Temperature (1) Continuous white light and 20°C n = 5r x 13s n = 4r x 9s n.a. n = 4r x 9s (2) Continuous white light and 30ºC n = 5r x 13s n = 4r x 9s n.a. n = 4r x 9s (3) Alternating between light and tempera- ture n = 5r x 13s n = 4r x 9s n = 3r x 7s n = 4r x 9s Experiment 4: Light Treatments (1) Continuous white light n = 4r x 13s n = 4r x 8 n = 3r x 7s n = 4r x 13s (2) Darkness n = 4r x 13s n = 4r x 8 n = 3r x 7s n = 4r x 13s (3) Continuous Red light n = 4r x 13s n = 4r x 8 n = 3r x 7s n = 4r x 13s (4) Continuous Far Red light n = 4r x 13s n = 4r x 8 n = 3r x 7s n = 4r x 13s (5) Combination of Red and Far Red light n = 4r x 13s n = 4r x 8 n.a. n = 4r x 13s 2014] 241 LATIFAH et al.: A physiological approach to conservation of four palm species to allow water absorption, because in nature the germination of seeds with a hard seed coat that is impermeable to water is stimulated when the seed coats are cracked or softened gradually by abrasion against soil and rocks (Bewley & Black, 1994). The general design of the experiments is presented in Table 3, including the number of replicates and seeds used in each experimental unit, which varied according to the number of seeds available. Species were not analysed as treatments as the experiments on each species were not conducted at the same time due to differences in seed availability; however, general comparisons between species were possible. The fully-imbibed seeds were sown between two water-saturated 9 cm-filter papers in petri dishes (ISTA, 1985). All treated seeds were then placed in the temperature-controlled room. As scarification could help water intrusion, based on previous work on palm seed imbibition (Latifah, 2004), all seeds were scarified in experiments 3 and 4, and germina- tion was monitored. Variables observed were: Water mass and time to full imbibition The amount (g) of water imbibed was determined per 10 seeds, because the mass of one seed was too small (Table 2). Water mass is the quantitative variable in the imbibition process and is described as the amount of water imbibed across the dor- mancy breaking treatments during the imbibition period. The imbibition period was determined as the time when there was no further change in the amount of water absorbed. Germination The quantitative variable in the seed germination experiments is ‘germination’ expressed as a per- centage. The formula is as follows: Germination = n/N x 100% (n = number of seeds that germinated, N = total seeds per experimental unit) Germination was defined by the appearance and elongation of the cotyledonary tubule in A. austral- asica and L. ramsayi var. tuckeri. In C. australis and H. wendlandiana, germination was defined as the appearance and elongation of the plumule. The number of seeds germinated was recorded regularly and then they were removed. After seeds ceased germinating, the viability of ungerminated seeds was tested using tetrazolium. The presence of vi- able ungerminated seeds indicated seed dormancy. Viability was calculated as: Viability= (n1+n2)/N x 100% (n1= number of seeds that germinated, n2= number of seeds that were viable according to the tetra- zolium test, N = total seeds per experimental unit). Viability was used to help interpret the germination results. The viability of the seeds was based on the staining pattern in embryos and endosperms (Bhodthipuks et al., 1996; ISTA, 1985). Since no information is available on palm seeds, this determi- nation was based on viability tests done on Ora- niopsis appendiculata and Arenga microcarpa (Latifah, 2004). As palm seeds have a hard en- dosperm and seed coat, the palm seed is determined as viable when the embryo is red and endosperm is red or more than 50% red. Time to first germination and final germination The rate of germination can be determined by several measures such as time to first germination, time to final germination, T50, coefficient rate of germination or coefficient rate of germination simultaneity (Bewley & Black, 1994; ISTA, 1985). However, as the palm seeds germinated slowly, time to first germination and time to final germina- tion were used to assess the germination rate. Time to first germination was the time (days after sowing) when the first seed germinated in each replicate. Time to final germination was time to the last signs of seed germination. The differences in the amount of water imbibed across the treatments over time and the differences of the time effect were analysed by repeated meas- ures ANOVA. Germination and time to final germination were analyzed by one-way analysis of variance (ANOVA) to determine if there were significant treatment effects. When the data were not normally distributed, the Kruskal Wallis Non Parametric Test was applied using SPSS version 17.0 package was used for all the analyses. RESULTS & DISCUSSION Arenga australasica belongs to the sub family Arecoideae, and all members of this group have a very thick seed coat (Uhl & Dransfield, 1987) (Fig. 1). The seed coats of A. australasica seeds slowed their imbibition (Fig. 2A; Table 4). The fully- imbibed scarified or heated seeds tended to have higher germination (82-88%) within 185-200 days (began to germinate after 40-42 days) than un- treated seeds (68%) within 167 days (began to germinate after 39 days). Hence fewer untreated seeds germinated, but they germinated faster. This REINWARDTIA 242 [VOL.14 Fig.2. Amount of water absorbed per 10 seeds (mean ± SEM) and time for imbibition of A). A. australasica, B). C. australis, C). H. wendlandiana and D). L. ramsayi in response to different dormancy breaking techniques: no treatment (□), scarification (Δ), moist heat 50°C (○). Samples with the same upper (treatment effect) or lower case letters (time) are not significantly different (p > 0.05); Rate*Time=interaction. Fig. 3. Germination and times to first and final germination (mean ± SEM) of (A-C) A. australasica, (D-F) C. australis and (G-I) L. ramsayi in response to different dormancy breaking techniques: no treatment ( ), scarification ( ) and moist heat 50 ºC ( ). Secondary histograms ( ) for L. ramsayi show seed viability (%) after adding the viable ungerminated seeds to the number of germinated seeds. F(2,9)=3.26,p=0.09), log transformed A Arenga australasica B Arenga australasica C 2014] 243 LATIFAH et al.: A physiological approach to conservation of four palm species is a high total germination compared to that found in previous studies of Arenga species. In one study, seeds of A. pinnata and A. wightii yielded only 27% and 4% germination and germinated after 27 and 3 days respectively, without any treat- ments (Rees in Koebernik, 1971). Dormancy of A. pinnata seeds was broken by acid or mechanical scarification (using a file) treatments after 30-365 days; a few samples germinated after 730 days (Mujahidin et al., 2003). A. microcarpa seeds gave 74-86% germination after warm stratification for 2-6 weeks at 27-40°C (Odetola, 1987). A. engleri, A. microcarpa, A. obtusifolia, A. pinnata, A. tremula, A. undulatifolia and A. wightii broke dormancy after 83-626 days without any treat- ments (Koebernik, 1971). In another study, scari- fied seeds of A. microcarpa had 70% germination after 63-308 days (Latifah, 2004). On the other hand, the seeds of A. australasica had 90% germi- nation within 19 days (ISTA, 2006); there is no information about the treatment, the source and the age of the seeds used in this study. The varying results above suggests that Arenga spp., including A. australasica, may have secondary dormancy (Baskin & Baskin, 2001; Fenner & Thompson, 2005; Vleeshouwers et al., 1995), which refers to seeds that are able to germinate after shedding from their parent trees when conditions are suitable for germination and seedling establish- ment; but the seeds can also lie dormant during seasons that are unfavourable for germination and establishment. Imbibition of C. australis seeds was also slowed by the seed coat (Fig. 2B; Table 4). The properties of the sarcotesta may cause water impermeability according to Bass (1979). No previous research on C. australis could be found; however, the literature contains some information for other Calamus species. In the current study, 78% of untreated seeds (sarcotesta removed (Fig. 1) germinated after 10 days (but within 37 days) (Fig. 3.D-F). In comparison, C. peregrinus showed 91% germina- tion within 12-35 days with the sarcotesta removed (Vangkualong, 1984). In the current study, 70% of C. australis seeds, which had been scarified or heated, germinated after 10-13 days (within 64-71 days); but, this was not significantly different to untreated seeds. Mohd et al. (1994) found increased germination of C. latifolius after heating at 40°C for 48 hours. The current study and the previous research by the other authors did not examine germination without removal of the sar- cotesta. Similarly, the seed coat of H. wendlandiana slowed imbibition (Fig. 2C; Table 4). As in A. aus- tralasica, H. wendlandiana also belongs to sub family Arecoideae whose members have a very thick seed coat (Uhl & Dransfield, 1987) covered by a fibrous sheath (Essig & Hernandez, 2002) (Fig. 1). Untreated seeds (the fibrous sheath was not removed) of H. wendlandiana gave 43% ger- mination after 102 days. The scarified and heated seeds did not germinate. Lawie (2007) found that scarified seeds of H. wendlandiana took 365 days for the first germination, but does not describe the percentage that germinated. Another worker found that fresh seeds without any treatment germinated within 90-180 days (Cronin, 1989). In the present study, scarification and moist heat did not promote germination. The seeds may require certain light conditions for germination (Fig. 6; Table 4). Imbibition was slowed by the seed coat of L. ramsayi (Fig. 2D; Table 4), as in the other three species. L. ramsayi belongs to sub family Cory- phoideae in which many members have very thick seed coats (Uhl & Dransfield, 1987). The seeds of this species also absorbed more water (0.08 mg/ seed) when fully imbibed after 48 hours than the other study species despite the seed weight being lighter than A. australasica and C. australis. In the current study, scarification gave 78% germina- tion after 101 days (within 216 days), which was higher than untreated (60%, after 133 days, within 211 days) and heated seeds (58%, after 109 days, within 199 days). Moreover, some of the ungermi- nated seeds were still viable, resulting in 71% (no treatment), 80% (scarification) and 77% (moist heat) viability. A previous study found that L. ramsayi var. ramsayi germinated slowly and erratically in the nursery as well as in rainforests. In the nursery, the germination of L. ramsayi var. tuckeri seeds took a year (Diy pers. comm., January 7th 2009). Previous research on other species of Licuala found L. elegans, L. gracilis, L. lauterbachii, L. muelleri, L. peltata and L. spinosa took 53-475 days to germinate without any treat- ments (Koebernik, 1971). These results suggest that seed coat-imposed dormancy is present and the endocarp that covers the seed coat may inhibit imbibition and germinationin L. ramsayi seeds. A. australasica seed germination was inhibited by far red light (Fig. 5A; Table 4). This suggests that canopy gaps created by cyclonic disturbances may induce seed germination in this species since: (1) this result was not significantly different from that in white light and (2) the far red and red-far red combination resulted in lower germination, 14% (72 days first germination) and 38% (130 days first germination). REINWARDTIA 244 [VOL.14 All of the C. australis seeds germinated under all light treatments (Table 4). This indicates that canopy gaps were also favourable for germination of C. australis. The germination of this species was not affected significantly by light condition. No previous research of this nature has been done on C. australis. No Hydriastele wendlandiana seeds germinated under far red light, while most germinated under darkness or red light (Fig. 6A; Table 4). Under white light only 43% untreated seeds germinated after 102 days (Experiment 2: Dormancy Breaking Treatments). The seed germination was inhibited by far red light. No previous research on effects of light on germination of this species could be found in the literature. However, these results suggest that untreated seeds germinated slowly under continuous white light and scarified seeds under white light did not germinate; although the scarified seeds germinated under darkness or red light. Similarly, most seeds of L. ramsayi germinated under red and white light and were inhibited by far red light (Fig. 5F; Table 4). This suggests that germination of L. ramsayi seeds may be favoured by canopy gaps. No similar studies could be found for L. ramsayi in the literature. Seed germination of the four palm species also occurred in darkness (Fig. 5-6; Table 4). The phytochrome system is involved in seed germina- tion in darkness, the seeds may have sufficient Pfr which is an active form that can promote germina- tion in darkness (Bewley & Black, 1994b). No research in this area has been done in A. austral- asica. However, in another Arenga species, A. mi- crocarpa, darkness resulted in low germination when the seeds were placed inside 500 gauge polyethylene bags and exposed to continuous Fig. 4. Germination and times to first and final germination (mean ± SEM) of (A-C) A. australasica, (D-F) C. australis and (G-I) L. ramsayi in response to different temperature treatments and diel fluctuation of light and temperature: 20 ºC full light ( ), 30 ºC full light ( ) and 20/30 ºC dark/light ( ). Samples with the same lower case letters are not significantly different (p=0.05). Secondary histograms ( ) for L. ramsayi show seed viability (%) after adding the viable ungerminated seeds to the number of germinated seeds. 2014] 245 LATIFAH et al.: A physiological approach to conservation of four palm species white light or continuous darkness (Odetola, 1987). The germination of the palms studied varied in response to different temperatures (20 and 30°C) and diel light and temperature fluctuations. The germination of A. australasica, C. australis and L. ramsayi was lower at 20°C, compared to 30°C. The effects of diel fluctuations were not significant in A. australasica and C. australis compared to L. ramsayi, which germinated best under the fluctuat- ing treatment. In contrast, diel fluctuations inhib- ited the germination of H. wendlandiana seeds. A review of the literature found no similar research on palms. A non-palm and invasive tropical plant, Mikania micrantha, germinated rapidly in 25, 30 15/30 °C (night/day) and an ambient temperature (24-32°C) suggesting it is widely adapted to a range of natural conditions (Yang et al., 2005). Temperature and light fluctuations did not signifi- cantly increase the R:FR (Red:Far Red light) ratios and the transformation of Prto Pfr is influenced by temperature (Pr is a physiologically inactive form of phytochrome activated by Red light and transformed into Pfr, an active form, that canbreak seed dormancy (Atwell et al., 1999)). CONCLUSION The hard seed coats of A. australasica, C. aus- tralis, H. wendlandiana and L. ramsayi slowed imbibition. Scarified seeds germinated best in A. australasica, C. australis and L. ramsayi. The ger- mination of all seeds was inhibited by far red light. The red light requirement suggests that canopy gaps created by cyclonic disturbances could promote their germination. This implies that in disturbed rainforests, the dispersal agents may play important roles in promoting germination of some palms by scarifying through the frugivore gut passage and specific placement in canopy gaps. ACKNOWLEDGEMENTS I acknowledge Dr. John Dowe, A/Prof. Betsy Jackes, A/Prof. Ross Coventry, Christopher Gardiner and Dr. Fig. 5. Germination and times to first and final germination (mean ± SEM) of (A-C) A. australasica, (D-E) C. australis and (F-H) L. ramsayi in response to different light treatments: continuous white light ( ), darkness ( ), continuous red light ( ), continuous far red light ( ). Samples with the same lower case letters are not significantly different (p=0.05). Secondary histograms ( ) for L. ramsayi show seed viability (%) after adding the viable ungerminated seeds to the number of germinated seeds. REINWARDTIA 246 [VOL.14 Leone Bielig for their intellectual support. 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Sin. 46: 293−299. http://www.seedtest.org/upload/cms/user/PresentationoftheForestTreeandShrubSeedCommittee.pdf http://www.seedtest.org/upload/cms/user/PresentationoftheForestTreeandShrubSeedCommittee.pdf http://www.iucnredlist.org http://www.iucnredlist.org REINWARDTIA 248 [VOL.14 INSTRUCTION TO AUTHORS Scope. Reinwardtia is a scientific irregular journal on plant taxonomy, plant ecology and ethnobotany published in December. Manuscript intended for a publication should be written in English. Titles. Titles should be brief, informative and followed by author's name and mailing address in one- paragraphed. Abstract. English abstract followed by Indonesian abstract of not more than 250 words. Keywords should be given below each abstract. Manuscript. Manuscript is original paper and represent an article which has not been published in any other journal or proceedings. 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Phylogenetic position and generic differentiation of Epithemateae (Gesneriaceae) inferred from plastid DNA sequence data. American J. Bot. 90: 321-329. Proceedings :TEMU, S. T. 1995. Peranan tumbuhan dan ternak dalam upacara adat "Djoka Dju" pada suku Lio, Ende, Flores, Nusa Tenggara Timur. In: NASUTION, E. (Ed.). Presiding Seminar dan Lokakarya Nasional Etnobotani II. LIP1 & Perpustakaan Nasional: 263-268. (In Indonesian). SIMBOLON, H. & MIRMANTO, E. 2000. Checklist of plant species in the peat swamp forests of Central Kalimantan, Indonesia. In: IWAKUMA et al. (Eds.) Proceedings of the International Symposium on: Tropical Peatlands. Pp. 179-190. Book : RIDLEY, H. N. 1923. Flora of the Malay Peninsula 2. L. Reeve & Co. Ltd, London. Part of Book : BENTHAM, G. 1876. Gesneriaceae. In: BENTHAM, G. & HOOKER, J. D. Genera plantarum 2. Lovell Reeve & Co., London. Pp. 990-1025. Thesis : BAIRD, L. 2002. A Grammar of Keo: An Austronesian language of East Nusantara. 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Author signature (s) Date Name MUHAMMAD EFFENDI, TATIK CHIKMAWATI & DEDY DARNAEDI. New cytotypes of Pteris ensiformis var. victoria from Indonesia 133 SUZANA SABRAN, REUBEN NILUS, JOAN T. PEREIRA & JOHN BAPTIST SUGAU. Contribution of the heart of Borneo (HoB) initiative towards botanical exploration in Sabah, Malaysia 137 WENNI SETYO LESTARI, BAYU ADJIE, TASSANAI JARUWATANAPHAN, YASUYUKI WATANO & MADE PHAR- MAWATI. Molecular phylogeny of maidenhair fern genus Adiantum (Pteridaceae) from Lesser Sunda Islands, Indonesia based on Rbcl and Trnl-f 143 ELIZABETH A. WIDJAJA & DANIEL POTTER. Floristic study of Mekongga Protected Forest: towards establishment of the Mekongga National Park 157 YESSI SANTIKA, EKA FATMAWATI TIHURUA & TEGUH TRIONO. Comparative leaves anatomy of Pandanus, Freycinetia and Sararanga (Pandanaceae) and their diagnostic value 163 SUHARDJONO PRAWIROATMODJO & KUSWATA KARTAWINATA. Floristic diversity and structural characteristics of mangrove forest of Raj a Ampat, West Papua, Indonesia 171 IAN M. TURNER. A new combination in Orophea (Annonaceae) for Uvaria nitida Roxb. ex G. Don 181 IVAN S AVINOV. Taxonomic revision of Asian genus Glyptopetalum Thwaites (Celastraceae R. Br.) 183 YUSI ROSALINA, NISYAWATL ERWIN NURDIN, JATNA SUPRIATNA & KUSWATA KARTAWINATA. Floristic compo- sition and structure of a peat swamp forest in the conservation area of the PT National Sago Prima, Selat Panjang, Riau, Indone- sia 193 IMAN HID AY AT & JAMJAN MEEBOON. Cercospora brunfelsiicola (Fungi, Mycosphaerellaceae), a new tropical Cercosporoid fungus on Brunfelsia uniflora 211 MAX VAN BALGOOY & ELIZABETH A. WIDJAJA. Flora of Bali: a provisional checklist 219 EKA FATMAWATI TIHURUA & INA ERLINAWATI. Leaf anatomy of Pandanus spp. (Pandanceae) from Sebangau and Bukit Baka-Bukit Raya National Park, Kalimantan, Indonesia 223 JULIA SANG & RUTH KIEW. Diversity of Begonia (Begoniaceae) in Borneo - How many species are there? 23 3 DIAN LATIFAH, ROBERT A. CONGDON & JOSEPH A. HOLTUM. A Physiological approach to conservation of four palm species: Arenga australasica, Calamus australis, Hydriastele wendlandiana saALicuala ramsayi 237 REINWARDTIA Vol. 14. No. 1.2014 CONTENTS Page ABDULROKHMAN KARTONEGORO & DANIEL POTTER. The Gesneriaceae of Sulawesi VI: the species from Mekongga Mts. with a new species of Cyrtandra described 1 LIM CHUNG LU & RUTH KIEW. Codonoboea (Gesneriaceae) sections in Peninsular Malaysia 13 WISNU H. ARDI, YAYAN W. C. KUSUMA, CARL E. LEWIS, ROSNIATI A. RISNA, HARRY WIRIADINATA, MELISSA E. ABDO & DANIEL C. THOMAS. Studies on Begonia (Begoniaceae) of the Molucca Islands I: Two new species from Halmahera, Indonesia, and an updated description of Begonia holosericea 19 YUZAMMI, JOKO R. WITONO & WILBERT L. A. HETTERSCHEID. Conservation status of Amorphophallus discophorus Backer & Alderw. (Araceae) in Java, Indonesia 27 MOHAMMAD F. ROYYANI & JOENI S. RAHAJOE. Behind the sacred tree: local people and their natural resources sustainabil- ity 35 FIFI GUS DWIYANTI, KOICHI KAMIYA & KO HARADA. Phylogeographic structure of the commercially important tropical tree species, Dryobalanops aromatica Gaertn. F. (Dipterocarpaceae) revealed by microsatellite markers 43 SACHIKO NISHIDA & HENK VAN DER WERFF. Do cuticle characters support the recognition of Alseodaphne, Nothaphoebe and Dehaasia as distinct genera? 53 NURUL AMAL LATIFF, RAHAYU SUKMARIA SUKRI & FAIZAH METALI. Nepenthes diversity and abundance in five habi- tats in Brunei Damssalam 67 NURUL HAZLINA ZATNI & RAHAYU SUKMARIA SUKRI. The diversity and abundance of ground herbs in lowland mixed Dipterocarp forest and heath forest in Brunei Darussalam 73 MUHAMMAD AMIRUL AIMAN AHMAD JUHARI, NORATNI TALIP, CHE NURUL ATNI CHE AMRI & MOHAMAD RUZI ABDUL RAHMAN. Trichomes morphology of petals in some species of Acanthaceae 79 DIAN ROSLEINE, EIZI SUZUKI, ATIH SUNDAWIATI, WARDI SEPTIANA & DESY EKAWATI. The effect of land use history on natural forest rehabilitation at corridor area of Gunung Halimun Salak National Park, West Java, Indonesia 85 JULIUS KULIP. The Ethnobotany of the Dusun people in Tikolod village, Tambunan district, Sabah, Malaysia 101 PETER O'BYRNE. On the evolution of Dipodium R. Br 123 Reinwardtia is a LIPI accredited Journal (517/AU2/P2MI-LIPI/04/2013) Herbarium Bogoriense Botany Division Research Center for Biology - Indonesian Institute of Sciences Cibinong Science Center Jln. Raya Jakarta - Bogor, Km 46 Cibinong 16911, P.O. Box 25 Cibinong Indonesia barudepan 421-609-2-PB belakangbaru img577_Page_1 img577_Page_2 img577_Page_3 img577_Page_4