REINWARDTIA_13_1_291209 today REINWARDTIA Vol 13, Part 1, pp: 45 − 67 45 EXPLORATION OF HIGH ELEVATION LIANA COLONIES ON MT. SLAMET, CENTRAL JAVA, INDONESIA Received June 8, 2009; accepted June 16, 2009 W.S. HOOVER New England Tropical Conservatory (NETC), BCIC, 940 Walter Street, North Bennington, VT 05257. USA. Email: netrop@sover.net DEDEN GIRMANSYAH Herbarium Bogoriense, Botany Division, Research Centre for Biology-LIPI, Jl. Raya Bogor-Jakarta km 46, Cibinong 16911, Indonesia. Email: Deden_bo@yahoo.com HARRY WIRIADINATA Herbarium Bogoriense, Botany Division, Research Centre for Biology-LIPI, Jl. Raya Bogor-Jakarta km 46, Cibinong 16911, Indonesia. Email: Harry_wiria@yahoo.com JAMES M. HUNTER New England Tropical Conservatory (NETC), BCIC, 940 Walter Street, North Bennington, VT 05257. USA. Email: info@netrop.org ABSTRACT HOOVER, WS., GIRMANSYAH, D., WIRIADINATA, H. & HUNTER, J. M. 2009. Exploration of High elevation liana colonies on Mt. Slamet, Central Java, Indonesia. Reinwardtia 13(1): 45–67. — One hundred forty–five individual lianas were distributed on 2 East facing ridges on the second highest mountain on Java, Mt. Slamet (3418 m.), Central Java, Indonesia. Twenty one colonies were observed on small flat areas on ridges. The liana species observed include: Embelia pergamacea, Toddalia asiatica, Elaeagnus latifolia, Schefflera lucida, Vaccinium laurifolium and Lonicera javanica. Diameter of each liana was measured and liana density/flat area calculated. Floristic collecting was under- taken within the elevational gradient of liana distribution. Data suggest an ecotone transition from lower to upper mon- tane forest is observed between 2200 and 2300 m, though forest types are difficult to determine due to disturbance caused by fire at the upper elevations. Observing lianas at these unusuall high elevations with near pluvial rainfall, con- tradict established scientific theory concerning global distribution and abundance of lianas. Keyword: Liana, Rainfall, Elevation, Mt. Slamet, Central Java ABSTRAK HOOVER, W. S., GIRMANSYAH, D., WIRIADINATA, H. & HUNTER, J. M. Eksplorasi koloni liana dataran tinggi di G. Slamet, Jawa Tengah, Indonesia. Reinwardtia 13(1): 45–67. — Seratus empat puluh lima liana tersebar di 2 lokasi yang menghadap punggung bukit sebelah timur G. Slamet (3418 m, gunung kedua tertinggi di Jawa), Jawa tengah, Indonesia. Dua puluh satu koloni terekam pada area datar punggung tersebut. Jenis–jenis liana yang tercatat adalah: Embelia pergamacea, Toddalia asiatica, Elaeagnus latifolia, Schefflera lucida, Vaccinium laurifolium dan Lonicera javanica. Diameter tiap–tiap liana diukur dan kepadatan liana dihitung. Data menunjukan daerah transisi (ecotone) ter- dapat pada ketinggian 2200 sampai 2300 m dpl, walaupun tipe hutannya tidak dapat di identifikasi karena bekas ter- bakar. Hasil pengamatan liana pada daerah dengan curah hujan yang sangat ekstrim tinggi dan ketinggian yang tidak biasanya tersebut bertentangan dengan teori keilmuan tentang distribusi global dan kelimpahan liana. Kata Kunci: Liana, Curah hujan, Ketinggian, G. Slamet, Jawa tengah INTRODUCTION Botanists observing tropical forests and climbers in the 19 th and early 20 th centuries noted the abun- dance of lianas at lower elevations and a decrease of these plants with increasing elevation (Treub 1883, Schenk 1892–3, Davis and Richards 1933–34, and Richards 1936). Such observations on lianas came to define a physiognomic characteristic of lowland tropical forests (Webb 1959, Grubb et al. 1963, Grubb and Whitmore 1966, Ashton 1964, Richards 1964, Whitmore 1984, 1988). Croat (1978) goes so REINWARDTIA 46 [VOL.13 far as to say the presence of woody climbers is the single most important physiognomic character dis- tinguishing tropical from temperate forests. Ashton (2003) considers 800–1200 m. an ecotone transition between lowland and lower montane forest, empha- sizing that different mountains have different char- acteristics, and vegetation types along tropical mountains cannot be identified by single, standard elevational definitions. In considering the transition from upper montane to subalpine, Kartawinata (2005) suggests the boundary be considered 2300 m., differing from Steenis et al. (1972) report that 2400 m. be the boundary, with much historical de- bate about elevation zones and tropical mountain vegetation. Kartawinata’s (2005) paper may be con- sidered a benchmark for identifying botanical and ecological research needs in Indonesia, including lianas; this paper is one of the first specific contri- butions ever on lianas of Indonesia. At the time Putz and Mooney (1991) published The Biology of Vines, research on lianas was limited compared to other plant groups. Gentry (1991), from this book, likely provides the first global data base on distribution and evolution of climbers; his report based on 56 neotropical and 32 paleotropical sites, all of which are lowland except for 9 montane neotropical forests sites, which are only briefly mentioned. Since 1991, scientific literature on li- anas has expanded greatly, with many papers exam- ining eco–physiological characteristics of climbers. Longino (1986) may have been the first to observe a negative correlation between growth and rainfall, while Schnitzer’s (2005) paper summarizes current eco–physiological research and provides explana- tion for global patterns of liana abundance and dis- tribution based on 69 neo–tropical lowland sites; elevation is not a factor mentioned in his work. Most of the literature on lianas over the past years has focused on neo–tropics, with comparatively limited research from SE Asia, though a number of papers are noteworthy (see for example: Jacobs 1976, Putz 1984, 1985, 1987, Campbell & New- berry. 1993, Pinard & Putz., 1993, Caballe 1994, Babweteera 2000, Dattaraja & Sukumar, 2004 and Parthasarathny et al. 2004). Kartawinata (2005:126) goes so far as to say, “Studies on lianas and epi- phytes should be promoted further since to date they have been neglected”, in reference to Indonesia. In the Mountain Flora of Java, Steenis et al. (1972) illustrate 12 species of lianas and about 14 other vine species having woody characteristics, noting that Toddalia asiatica and Vaccinium lauri- folium are 2 species from this present study men- tioned by van Steenis et al. (1972). V. laurifolium is documented to occur as a shrub on the summit of Mt. Gede/Pangrango complex, outside of the plots (Sadili et al. 2009) as well as having been observed to occur epiphytically (Wiriadinata, pers. comm.). Lonicera javanica is noted to be a highly invasive vine with a cosmopolitan distribution, likely con- sidered a weed (Dillenburg et al. 1993). Mt. Slamet (3418 m, 7 14’23.8” S. lat.; 109 13’.8” E. long.) was selected for exploration be- cause it is the second highest mountain on Java (Fig. 1 and 2); reporting the highest precipitation on the island is from the town of Baturden on the South slope of Mt. Slamet, averaging about 7000 mm/year (Fig. 3, Berlage 1941 and Indonesian Meterological and Geophysical Institute) and 2 nd highest reported rainfall in Indonesia (Schmidt & Ferguson 1951). Rainfall is noted to have exceeded 10,000 mm/year at Baturden (Fig. 1). Early exploratory research on Mt. Slamet is ex- tensive though appears limited to primarily lowland tropical forest, likely due to the relative in- accessibility of upper elevations due to extensive forest cover. The first botanist to explore Mt. Slamet appears to be G.F.W. Junghuhn with explo- ration made primarily between 1940–1945 (Steenis –Kruseman 1950). The Cyclopedia of Malaysian Plant Collectors (Steenis–Kruseman 1950) lists the following botanists as having undertaken explora- tion of Mt. Slamet, together with many other loca- tions in Indonesia, with their concentrated years of exploration: S.H. Koorders 1893–1914, P. de Mon- chy 1897, M. Raciborski 1897–1902, F.A.F.C. Went 1898–1930, C.A. Backer 1907–1948, L.G. den Berger 1917–1926, H.J. Lam 1924–1925, J. Jeswiet 1926, Hagederon 1926, C.G.G. J. van Steenis 1928–1940, F. Verdoorn 1935, F.C. Dre- scher 1958. Recent expeditions and collecting trips to Mt. Slamet are recorded from Bogor Herbarium’s digi- tal library including: Sulistyani (1995), Arifiani (2001), Rahman (2001), Susana (2002), and Wiri- adinata (2004) NETC sponsored expedition involv- ing the present authors. This paper reports on liana colonies forming a component of upper elevation tropical forest along two east–facing ridges on Mt. Slamet, Central Java, Indonesia, indicating lianas, in rare instances, are not restricted to lowland tropical forests but can become abundant at high elevations. Further, Mt. Slamet’s liana colonies thrive under pluvial–like rainfall conditions, furthering the distinction of this habitat. These liana data from Mt. Slamet contradict much of the conventional thinking on climbers. METHODS General floristic collecting was undertaken along HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 47 Table1. Vine Species from Mt. Slamet Species/Family/Local Name No. of Individuals No. of Colony Elevation (m) Trunk diameter (cm) Range Mean Range Mean Embelia pergamacea 9 1 1935 –1935 1935 +/– 0 2.6 –3.3 3.0 +/– 0.2 A.D.C. ( Myrsinaceae ) Gang Toddalia asiatica (L.) 9 4 1951 –2357 2243 +/– 169 1.4 – 4.5 2.4 +/– 1.1 Lam. ( Rutaceae ). Kuwut Vaccinium laurifolium 64 13 1966 –2591 2401 +/– 75 1.6 – 7.6 3.4 +/– 1.5 var. laurifolium Miq. (Ericaceae), Kematus Elaeagnus latifolia L. 11 3 1981 –2149 2068 +/– 60 1.4 – 3.2 2.6 +/– 0.6 (Elaeagnaceae) Duren Schefflera lucida L. 12 3 1981– 2316 2283 +/– 97 1.5 – 4.9 2.9 +/– 1.1 (Araliaceae) Tanganan Lonicera javanica DC. 40 4 2499 –2591 2552 +/– 22 1.3 – 3.7 2.2 +/– 0.5 (Caprifoliaceae ) Blebur Table 2. Colony Size and Distribution on Mt. Slamet Ridges 1 and 2 Total Ele- vation No of Total Colony Size No of vines / square Mean Change (m) Colony No of Vines (square m) meter Diameter (cm) Ridge 1 lower ½ 655 321 15 8 96 (66%) 34 (35%) 1025.17 675.17 0.18 0.18 3.2 3.3 Upper ½ 290 7 62 (65%) 350.00 0.19 3.0 Ridge 2 61 6 49 ( 34%) 1268.50 0.12 4.4 the entire elevation gradient from lowland until high elevation and data of vine were collected follow standard methodology for censuring and measuring liana (Schnitzer et al. 2006 and Gerwing et al. 2006). Specimens were collected and put between old news paper, pored with alcohol 70% put into big plastic bag and sent to Herbarium Bogoriense for drying and identification. RESULTS Herbarium specimens collected from the 6 spe- cies of lianas were identified at Bogor Herbarium and data summarized in Table 1. A clear distinction is observed between the elevational distribution and abundance of the 4 “lower elevation” species Embe- lia pergamacea, Elaeagnus latifolia, Toddalia asi- atica and Schefflera lucida and the 2 “upper eleva- tion” species Vaccinium laurifolium and Lonicera japonica. The former 4 species are much less abun- dant, with 41 individuals (28%) and distributed between 1935 to 2357 m, than the latter 2 species, with 104 individuals (72%) and distributed between 1966 to 2591 m; only 9 individuals of Vaccinium are found at the “lower elevation” with 55 found at the “upper elevation”. REINWARDTIA 48 [VOL.13 1\13 Vaccinium laurifolium 9 100 2469 2.99 +/– 1.71 0.090 1\14 Lonicera javanica 10 100 2499 2.36 +/– 60.00 0.100 1\15 Lonicera javanica 3 25 2591 2.86 +/– 0.28 0.120 2\16 Vaccinium laurifolium 2 10.5 2591 5.33 +/– 0.11 0.190 2\17 Vaccinium laurifolium 7 130 2560 3.50 +/– 1.50 0.054 2\18 Vaccinium laurifolium 5 930 2560 4.36 +/– 2.23 0.005 Lonicera javanica 21 930 2560 2.02 +/– 0.63 0.023 2\19 Vaccinium laurifolium 7 150 2545 3.34 +/– 1.13 0.047 2\20 Vaccinium laurifolium 1 33 2530 5.73 +/– 0.00 0.030 2\21 Lonicera javanica 6 15 2491 1.99 +/– 0.09 0.400 Table 2 indicates colony size and distribution of liana between both ridges. Ninety–six colonies of lianas (66%) are distributed on Ridge 1, while Ridge 2 has 49 colonies of lianas (34%). The eleva- tion gradient of Ridge 1 is 655 m., while Ridge 2 has 61 m. of gradient change. Observing colony size of available habitat for the liana, a distinction is noted between the 2 ridges. Ridge 1 has a total of 1,025 m 2 of flat colony surface area, or colony size; flat is a relative term in so far as we are on a steep mountain ridge. Nonetheless, the lianas were observed restricted to these flat areas since these locations were the only places where the lianas grew, and not on steep areas. Ridge 2 has a total of 1,268 m2 of flat colony habitat for the lianas, repre- senting roughly 20% more habitat surface than Ridge\ Colony No. Species No. of Individuals Colony size (Square Meters) Elevation ( meters ) Mean Diameter ( cm ) No of lianas/ square meter 1\1 Embellia pergamacea 9 17.2 1935 2.98 +/– 0.23 0.523 1\2 Toddalia asiatica 2 2.25 1951 1.51 +/– 0.11 0.889 1\3 Vaccinium laurifolium 2 16.72 1966 4.93 +/– 1.58 0.120 1\4 Elaeagnus latifolium 2 59.5 1981 2.63 +/– 0.34 0.034 1\5 1 59.5 1981 4.93 +/– 0.00 0.017 1\6 Elaeagnus latifolium 6 20 2057 2.68 +/– 0.50 0.300 1\7 Vaccinium laurifolium 6 400 2149 2.55 +/– 0.65 0.015 Elaeagnus latifolium 3 400 2149 1.80 +/– 0.33 0.008 1\8 Vaccinium laurifolium 1 100 2256 4.62 +/– 0.00 0.010 Schefflera lucida 1 100 2256 4.77 +/– 0.00 0.010 Toddalia asiatica 1 100 2256 3.18 +/– 0.00 0.010 1\9 Toddalia asiatica 1 50 2301 4.46 +/– 0.00 0.020 Vaccinium laurifolium 7 50 2301 2.41 +/– 0.65 0.140 1\10 Vaccinium laurifolium 8 25 2316 2.42 +/– 0.17 0.320 10 25 2316 2.45 +/– 0.73 0.400 1\11 Vaccinium laurifolium 3 25 2347 3.87 +/– 1.94 0.120 Toddalia asiatica 5 25 2347 2.1 +/– 0.74 0.200 1\12 Vaccinium laurifolium 6 25 2408 4.03 +/– 1.70 0.240 Table 3. Vine Species Diversity/ Colony and Individual Colony Data HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 49 Ridge 1, even with only 61 m. of elevation change, and being much shorter in distance. Overall liana density, measured by number of vines/ha is signifi- cantly different between the two ridges: Ridge 1 has a mean density of 1800 lianas/ha., while Ridge 2 is lower with a mean of 1200 lianas/ha. The opposite situation is observed for mean liana diameter: Ridge 1 having a mean liana diameter of 3.2 cm. and Ridge 2 having a mean of 4.4 cm. From Table 2, focusing on Ridge 1 with its 655 m. of elevational gradient presents its own set of interesting data. Most notable is greater abundance of lianas at the upper elevations: 34 lianas (35%) observed on the lower half of Ridge 1 and 62 lianas (65%) observed at the upper half of the ridge. The elevational gradient is not exactly divided in half for a “lower half” and an “upper half” because of plot locations, but the elevational dividing point is between Plot 8 (2,256 m) and 9 (2,301 m). The ele- vational change for the lower half is 321 m. and for the upper half is 290 m. Colony size for habitat is significantly different between “lower” and “upper” halves: the lower half of the gradient has 675 m 2 of surface colony area while the upper colony surface area is 350 m 2 . Mean diameter is different between the lower and upper halves of the ridge also: the lower with a mean diameter of 3.3 cm. and the up- per with a mean of 3.0 cm. Species diversity on colonies is of interest (Table 3). The lowest 6 colonies, from 1,935 m to 2,057 m on Ridge 1, are each consists of a single species. Embelia pergamacea, with 9 individuals, is observed only on colony number 1 and is not observed anywhere else on the 2 ridges. A similar situation for single species occupancy occurs at the highest most elevations also, where single species mostly dominate individual colonies: on Ridge 1, colonies 12–15 are each occupied by Vaccinium laurifolium or by Lonicera japonica. On Ridge 2, colonies 16–21, the same species occur with one or the other exclusively, except on the largest colony observed (930 m 2 ), number 18, which includes both species, though dominated heavily by Vaccinium. The middle elevation colonies on Ridge 1 are of interest from a multiple species standpoint: Colo- nies 7–11, ranging from 2,149 m to 2,347 m are occupied by 2 species on each flat area, except for colony number 8 (2,256 m), the only plot observed to have 3 species, V. laurifolium, Schefflera lucida and Toddalia asistica, with just one individual of each species. Colony number 8 (2256 m) is 100 m 2 of flat sur- face and is of special interest. Aside from its uniqueness with 3 species, if ranked, colony number 8 has one of the lowest liana densities (.01) of all colonies on Mt. Slamet, is the middle–most colony on Ridge 1, and the diameters of each liana rank among the largest measured: the Schefflera ranks 4th largest, Vaccinium ranks 5th and the Toddalia is 12 th , out of a total of 145 lianas colonies. Such cir- cumstances suggest explanation other than just co- incidence. The uniqueness of colony 8 suggests there may be other ecological or climatic charac- teristics of interest, at or near, this elevation, to be discussed below. Of the 21 colonies occupied by these lianas on Mt. Slamet, 14 (66%) are dominated by a single species, 6 (29%) diversified with 2 spe- cies each and colony number 8 (5%) with 3 species. Table 4 is a list of the general flora collected within the elevational range of the lianas (1935– 2591 m) on Mt. Slamet and compares the elevations of these collections with those of other sites throughout Java. Including the lianas, 19 species of general flora were collected. Other species were collected below and above the lianas, though are not included in this report. Interestingly, 8 species col- lected on Mt. Slamet from our expedition achieve elevational records for Java including: Saurauia microphylla, Schefflera aomatica, observed once, and S. lucida, a Mt. Slamet liana abundantly ob- served for this study, Vaccinium laurifolium, from this study, Peperomia tomentosa, only the 2nd speci- men collected on Java, Smilax odoratissima, Elatos- temma strigosum and Antrophyum reticulatum. Most of the general flora collected are common to Java, as indicated by the abundant collections from the island. When the Mt. Slamet lianas are exam- ined by rarity on Java, Vaccinium laurifolium var. laurifolium has only been collected 8 other times on the island, and is the most abundant vine observed on Mt. Slamet (64 individuals); this species ob- served as an epiphyte, also (Wiriadinata, pers. comm.). Elaeagnus latifolia has only been collected 14 times on Java, with 11 individuals on Mt. Slamet (Abdulhadi et al. 1998, Kartawinata 2005). DISCUSSION Mt.Slamet’s high elevation liana colonies appear distinct as a tropical ecology observation. This dis- cussion will examine these liana colonies in terms of a number of parameters including: forest type, elevation, taxonomic diversity, precipitation, pio- neering capability of lianas and fire as a cause for disturbance. How each of these parameters is re- lated to reported observations in tropical ecology will be discussed, indicating the distinction of Mt. Slamet’s upper elevation forest. Forest Type A central observation regarding Mt. Slamet’s upper elevation forest is its secondary nature, based on disturbance caused by fire, more of which will REINWARDTIA 50 [VOL.13 No Family, Genus, Species Collection from Java Our 2004 Mt. Slamet Collections Elev. record # Elev. Range Mean 1 Actinidiaceae Saurauia microphylla vriese 10 1400–1900 1500 1900&2500 X 2 Araliaceae Schefflera aromatica (Bl.) Harms Schefflera lucida L. 11 17 1400–1900 1300–2229 1782 1632 2000 2300 X X 3 Asteraceae Gynura aurantiaca DC. 31 600–2550 1560 2200 4 Caprifoliaceae Viburnum lutescens Bl. Lonicera javaniva DC. 53 31 150–3000 1000–3200 1112 1598 2500 2700 5 Cyperaceae Carex filicina Nees 20 1400–3020 2349 2500 6 Elaeagnaceae Elaeagnus latifolia L. 14 250–2000 1211 2300 7 Equisetaceae Equisetum debile Roxb. 65 100–2680 1317 2500 8 Ericaceae Vaccinium laurifolium var laurifolium Bl. 8 600–2300 1596 1500/2500/2900 X 9 Fagaceae Lithocarpus spicatus (Sm) Rehd et Wills 44 93–2403 1150 2600 10 Myrsinaceae Ardisia javanica DC. Embelia pergamacea DC. 87 20 93–3000 300–2400 2500 2100 2500 2100 11 Piperaceae Peperomia laevifolia Miq. Peperomia tomentosa Dietr. Piper caninum Bl. 91 1 96 200–2200 1642 25–2708 1425 1642 546 1700/1900 1900/2200 2000/2100/2700 X 12 Polypodiaceae Microsorium nigrescens Bl. 35 200–2000 1061 1900 13 Ranunculaceae Ranunculus blumei Steud Thalictrum javanicum Bl. 92 58 700–3300 1000–3300 2007 2739 1690/2500/2800/29 50 2500 14 Rutaceae Toddalia asiatica (L.) Lam. 47 93–2600 1719 2600 15 Saxifragaceae Dichroa febrifuga Lour. 91 500–3000 1530 1400/2200/2800 16 Smilacaceae Smilax odoratissima Bl. Smilax zeylanica L. 30 63 900–2200 500–2000 1642 626 1300/1900/3000 1900 X 17 Symplocaceae Symplocos cochinchinensis (Lour.) Moore 12 7 150–3020 1532 2600 18 Urticaceae Elatostemma strigosum (Bl.) Hassk. Pilea angulata Bl. Urtica bullata Bl. 9 38 12 970–2933 800–2600 1500–2600 1707 1460 2049 2200/2950 2000 2100 X 19 Vittariaceae Antrophyum reticulatum (Forst.)Kaulf. 64 5–1846 450 2000 X Tale 4. General Species List from Mt. Slamet HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 51 be said below. An observational gradient of distur- bance is evident along Ridge 1 from 2300 m. to the “burn–off zone” at 3,076 m. Lower elevational forest, estimated from 1850–2300 m., may be the richest montane forest we have observed in Indone- sia (Hoover et al. 2004), suggesting its disturbance is relatively limited. Above 2300 m, indications of more recent disturbance are observed, with the dis- tribution of lianas being a prime indication since total number of lianas increases dramatically after 2300 m (Fig. 9). It is important to attempt identification of the type of forest in which these Mt. Slamet lianas are growing, which is montane. Ashton (2003) points out it is often difficult to distinguish floristic, struc- tural and physiognomic elements separating lower montane from upper montane forest, thus describing an ecotone transition between these vegetation zones. Standard elevational benchmarks for forest transitions are unreliable because mountains have different heights and are subject to varying climatic conditions. The studies by Grubb et al. (1963) and Grubb and Whitmore (1966) in Ecuador, Whit- more’s (1984) studies of tropical mountains of Asia and throughout the tropics and (Whitmore 1998) all are classic studies of tropical forest zonation in rela- tion to elevation. More recent studies have continued to examine complex issues regarding vegetational zonation on SE Asian tropical moun- tains, with little agreement among scientists; see for example, but not limited to Osawa (1991), Kita- yama (1992), Gioda et al. (1993), Aiba and Kita- yama (1999), Beaman et al. (2001) and Kartawinata (2005). Nonetheless, Ashton (2003) makes a clear case for transitional/ecotone tropical forest zones that can be effectly applied in the field. Table 4 lists the general flora associated with the lianas of Mt. Slamet and compares the elevational ranges and means of these species with the same species collected throughout Java. It is predictable a number of floristic elevational records (8) would be achieved since Mt. Slamet is the second highest mountain on Java. Examining species associated with lianas suggests a floristic composition repre- sentative of lower montane forest on Java, also sug- gested by Ashton (pers. comm.), with elevational records achieved by these species simple because forest habitat extended by high elevation. The forest observed along the elevational gradient up to 2200 m. is likely lower montane with an ecotone transi- tion roughly occurring between this elevation and 2300 m.; within this elevation range occur a number of distinctions in the liana data: 1. Plot 8 (2256 m), being the only plot observed with 3 species of lianas, occurring in the middle of the gradient and concentrated with 3 of the largest diameter lianas observed. 2. The 4 “lower elevation” species do not exceed 2347 m, while the 2 “upper elevation” species are concentrated above 2301 m at colony num- ber 9, with only 9 Vaccinium occurring below 2300 m. 3. Fig. 9 indicate data at the 2201–2300 m eleva- tional range category are distinct since each of the parameters (total number of individuals, mean lianas/m 2 and mean trunk diameter are respectively lowest, one of the smallest and highest. The disturbed nature of the upper elevation forest complicates identification of forest type, be it lower or upper montane, and at what elevation forest tran- sition occurs is difficult to assess. However, the data indicate distinction around 2200–2300 m. It is of interest to note Kartawinata (2005) states, “the altitude of 2300 m is the meeting point of dominant species of upper montane forest and sub– alpine forest, hence the elevation of 2300 may be suggested as the boundary between montane zone and subalpine differing from the boundary at alti- tude 2400 m. previously defined by van Steenis et al (1972)”. Mt. Slamet is distinct from the Mt. Gede/ Pangrango complex since no vegetation exists on the top 400 m of Mt. Slamet. Fire has completely burned off the top of Mt. Slamet, while Mt. Gede/ Pangrango has experienced a limited recent fire his- tory. None–the–less, 2300 m certainly appears to be a critical elevation for SE Asian tropical forests. Regarding Mt. Slamet, the elevational range around 2200–2300 m. may to be an ecotone transition zone between lower and upper montane forest. Though not part of this study, it should be pointed out that Lonicera lianas were observed on the upper most sections of Ridge 2 at approximately 2700 m, but they appeared to be very young plants, with diameters of less than 10 mm. Evidence of fire was observed on the trunks of small trees. It was difficult to assess this forest as upper montane, due to the apparent disturbance caused by fire, but there still appeared to be enough distinguishing character- istics such as low stature and single canopy tree stratum, few shrubs, and microphyll leaf size to dis- tinguish this burned forest as upper montane. Elevation In itself, the presence of lianas on mountain slopes is not unusual. Steenis et al. (1972) report on 14 different species of lianas in the Mt. Flora of Java, with an additional number of climbers having woody characteristics. Most of Steenis et al. (1972) liana are observed at elevations closer to 1000 – 1500 m., much less than the Mt. Slamet site REINWARDTIA 52 [VOL.13 beginning at 1935 m and above. Gentry (1991) lists his 9 montane forest sites of the neotropics where lianas are present by elevation. The Pasochoa, Ecuador (3,010 m) site is higher than Mt. Slamet’s liana site, though this Ecuadorian site is based on only a 0.1 ha and the data extrapolated. All Gentry’s 9 montane sites are listed as a single elevation, whereas the Mt. Slamet site is along an elevational gradient from 1,935 – 2,591 m. Aside from this re- port and Gentry’s (1991), elevation is generally not a factor in the great volume of recent literature since Putz and Mooney (1991) published their “Biology of Vines”. In order to establish data standards, Schnitzer (2005), “…omitted forests that were clas- sified as subtropical, montane and premontane to reduce confounding the differences in latitude, ele- vation, and temperature among the sites.” A review of the last 18 years of liana literature indicates re- search is restricted to lowland tropical forest, pri- marily of the neotropics, where elevation is not a parameter examined. Therefore, the distinction with Mt. Slamet’s lianas is their colony structure distributed along a 655 m elevational gradient, with the greatest abundance of plants found at the higher elevations, particularily because of Vaccinium lauri- folium and Lonicera japonica. Taxonomic Diversity The 6 lianas observed at Mt. Slamet, Embelia pergamacea (Myrsinaceae), Elaeagnus latifolia (Elaeaganaceae), Toddalia asiatica (Rutaceae), Schefflera lucida (Araliaceae), Vaccinium lauri- folium (Ericaceae) and Lonicera japonica (Caprifoliaceae) represent families that are different from the limited Asian data sets available. Gentry’s (1991) 3 Asian sites were collected from lowland forest in Kalimantan, and would be expected to be different, with his 7 most common liana families being: Annonaceae, Apocynaceae, Fabaceae, Lo- ganiaceae, Rubiaceae, Connaraceae and Dil- leniaceae. Considering the taxonomic composition of the lianas at his 9 Neotropical upland Andean sites, only the Ericaceae is shared with the Mt. Slamet site, though he is only reporting the most common families, noting that lianas from many other families are observed. Of interest is that Gen- try (1991) points out that the Asteraceae is the most speciose Neotropical montane vine family and in general Andean montane forest climbers are domi- nated by a hemiepiphytic habit, as opposed to ground based woody lianas, like the 6 Mt. Slamet, species, further distinguishing this site. Precipitation Precipitation at Baturden, Central Java is ranked as the second highest measured in Indonesia, aver- aging over 7,000 mm/year for the last 93 years (Fig. 3a). (For interest’s sake the 5 highest measured rainfall sites in Indonesia are, by site, province, mean rainfall/year in mm, and the years data were collected (Berlage 1949 and Schmidt & Ferguson 1951): 1. Sungai Batoeng, W. Sumatra, 7752 mm, 1929–1936; 2. Tenjo, Batuaden, C. Java, 7069 mm, 1919–1941; 3. Tombo, Ond., C. Java, 6656 mm, 1889–1941; 4. Patoengkrinono, C. Java, 6649 mm, 1897–1941; 5. Pagilaran, C. Java, 6379 mm, 1896– 1941. There may well be wetter areas in Papua.) This is nearly a “Pluvial” level of rainfall by the Holdridge (1967) system for Neotropical Life Zones, referring to this system because there ap- pears to no eqivalent category for SE Asia. Pluvial Forest is benchmarked at 7400 mm/year and above by Gentry (1991). Though precipitation is not known specifically at the Mt. Slamet liana site, its adjacency to Baturaden suggests the site is very wet. (Appendix 2 shows old historical maps for rainfall stations and rainfall patterns at Baturaden (number 25) and Mt. Slamet, noting stream density on the south slope. Further, the next most adjacent sites to our Mt. Slamet liana site, besides Baturden being the closest, include sites due East, with each site number and mean rainfall: number 23J jelegong – 6012, number 23a Noesakambang – 5952 and number 24 Redjasari – 5565 mm Berlage 1949) For reference, Gentry (1991) only reports on 2 of his liana sites where rainfall exceeds Baturaden: Tutun- endo, Colombia with 9000 mm/year at an elevation of 90 m. and Mt. Cameroun, Cameroun with 8000 mm/year at 230 m. elevation, both lowland sites. No precipitation data are available for any of Gentry’s (1991) 9 montane forest sites. A review of available literature suggests liana abundance and distribution is negatively correlated with precipitation. The mechanistic explanation for global liana patterns of distribution offered by Schnitzer (2005) is based on a data set including 66 tropical forests: 24 dry forests, 31 moist forests and 11 wet forests. Schnitzer’s (2005) explanation for greater abundance and distribution of lianas in dry forest habitats involves the adaptive capability of lianas to exploit dry seasonal conditions through faster growth rates compared to trees, and a deep root system allowing plants to obtain necessary wa- ter. Research supporting Schnitzer’s (2005) overall theory is spread far and wide in the recent literature noting particularily, but not limited to: Longino (1986), who may have been the first to observe this negative correlation between liana abundance and precipitation, Gartner et al. (1990), Ewers et al. (1991), Opler et al. (1994), Philips et al. (2002), Restom and Nepstad (2004), and Kalacska et al. (2005). Thus, Mt. Slamet’s liana community distrib- uted over 655 m of elevational gradient in a “hyper– HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 53 wet”, high elevation forest appears distinctive for global tropical ecology. Pioneering Capability of Mt. Slamet Lianas and Fire as a Cause for Disturbance Noted throughout tropical literature, are lianas pioneering capacity to invade disturbed habitats, in reference to tree fall gaps, hurricanes, and logging sites (see for example: Putz et al. 1984, Balee and Campbell 1990, Fisher and Ewers 1991, Bab- weteera et al. 2000, Schnitzer 2005, Avalos et al. 2007. Additional tropical literature primarily from SE Asia describes the ecological effects of distur- bance caused by fire. See for example: Kartawinata (1993), Riswan (1982), Riswan and Kartawinata (1988, 1989, 1991), Sadili et al. (2009) describe the effects of fire on vegetation patterns on Mt. Gede Pangrango complex, West Java, Indonesia. Similarily, Mt. Slamet is a volcano and the ef- fects of fire were apparent on the trunks of the Aca- cia sp. and Anaphalis viscida at the subalpine zone, as well as at a section of upper montane forest at 2700 m. on Ridge 2, observed above the liana site. It is proposed that fire may be an environmental factor effecting the upper elevation forest on Mt. Slamet, and contributes directly to the presence of the lianas, due to their pioneering ability. Though the specific liana colonies showed no direct or immediate evidence of fire, this does not preclude the possibility that fire has burned areas along the ridges now occupied by lianas, thus disturbing the ecology of the forest and allowing lianas to invade with their rapid growth rates and form the colonies observed. The distribution of lianas on Mt. Slamet may be explained by the disturbance caused by fire, particularily, the greater abundance of Lonicera and Vaccinium at the upper elevations, as mentioned above. It is our hypothesis that fire has been a frequent factor influencing the upper elevational forest ecology of Mt. Slamet for much of its history, to a point where the steep upper section of the mountain on its South face, estimated at 800 or more meters of elevational gradient, is basically a “vine or liana forest”, not in the sense of Balee & Campbell (1990) or Perez–Salicrup (2001) Amazonian Liana Forests but a localized one. We hypothesize that by rotating from the steep South side of the mountain to the much less steep east side of the mountain, montane forest will become more prevelant, and thus exhibit a gradient in liana abundance and den- sity. Rotating further to the NE side of the moun- tain, where the present site has been documented, one observes even fewer lianas than the east side. In other words, the highest liana density and abun- dance has not been observed and documented, but is predicted for the South slope. The steep upper South slope has likely been effected not only by fire, but also by landslides due to being so steep, as Fig. 1. View of Mount Slamet showing upper section of western ridge from Baturaden REINWARDTIA 54 [VOL.13 can be observed in Fig. 2, thus causing further dis- turbance of this area and promoting the pioneering capability of lianas to invade and dominate the ecol- ogy. At this point, we plan a dry season observa- tional trip up the South slope to document photo- graphically whether the hypothesized liana commu- nity exists as a “Liana Forest” at a local scale. Climate Change It would be irresponsible not to mention the pos- sible effects of climate change on montane forest of Mt. Slamet, especially in view of Svenning & Con- dit (2008) commentary on the lack of research on climate change in the tropics. Perhaps climate change has contributed to a shift in montane forest vegetation on Mt. Slamet since this liana commu- nity appears unusual. Rainfall data from Baturaden (Fig. 3) suggest a decline in precipitation over the last 93 years, still leaving precipitation very high on Mt. Slamet’s South and East slopes. Such a shift in rainfall may be explained in a number of ways. 1. Overall global climate change is effecting rainfall patterns locally in Java. 2. Forest clearing; perhaps up through the 1950’s much of the lowland coastal area of Central Java was forested and by the early 1960’s this forest was destroyed. Philips et al. (2002) and Wright et al. (2004) have data suggesting sharp increases in liana abun- dance and productivity in neo–tropical forests, while Schnitzer (2005) acknowledges the possible effects of climate change on liana abundance and distribution. From the standpoint of elevation shifts of biota in the tropics, Colwell et al. (2008) present a new concept for approaching climate change in the tropics, with perhaps future consideration being given to applying this model to the upper elevation lianas on Mt. Slamet. Certainly, mankind’s assult on nature may produce consequential changes in vege- tation with perhaps more ecological circumstances like Mt. Slamet being observed in the future. Of interest to the history of tropical ecology and research on lianas, is the important role of Longino, sparked in part by Colwell et al. (2008) paper, in which Longino is one of the authors. As mentioned above, Longino (1986) paper in Biotropica may likely be the first research to document the negative correlation between lianas and precipitation; an ob- servation of great importance to global tropical ecology, witnessed by the great volume of post– 1991 literature concerning this observation. Of the many important post–1991 studies, Longino does not appear to have contributed much beyond the original, critical observation in 1986. Gentry (1991:14) does not make reference to this paper, though his data report, “It is probably not altogether coincidental that the neotropical site with the high- est liana density (Jauneche, Ecuador) occurs in an area with a marked dry season that is transitional between moist and dry forest.” What is the explana- tion for J.T. Longino’s highly influential, original observation on lianas being elaborated by other sci- entists who published much more?. Curiously, a similar question can be asked about why Dr. Enri- que Forero, Universidad National in Bogota, Co- lombia, was not a second author with Gentry on the many papers he published on work in Colombia, where Forero was his active partner and undertook Serang Jakarta Bandung G. Slamet Jogjakarta Semarang Surabaya Malang Banyuwangi Banten Prov. Jakarta Prov. W.Java C. Java Prov. E. Java Prov. Jogjakarta Prov. Fig. 2. Map of Java Island with Mount Slamet, showing major cities in Provinces HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 55 3000 4000 5000 6000 7000 8000 9000 10000 11000 19 16 19 18 19 20 19 22 19 24 19 26 19 28 19 30 19 32 19 34 19 36 19 38 19 40 19 42 19 44 19 46 19 48 19 50 19 52 19 54 19 56 19 58 19 60 19 62 19 64 19 66 19 68 19 70 19 72 19 74 19 76 19 78 19 80 19 82 19 84 19 86 19 88 19 90 19 92 19 94 19 96 19 98 20 00 20 02 20 04 20 06 20 08 0 100 200 300 400 500 600 700 800 900 1000 JA N F E B M A R A P R M E I JU N E JU L A U G S E P O K T N O P D E S Mean Monthly Rainfall # o f m m /M o n th Year Fig 3. Rainfall Data for Baturaden, Central Java ( Berlage 1949). a. Annual from 1916–2008 b. Monthly Mean Rainfall a. b. REINWARDTIA 56 [VOL.13 2 cm A B C D Fig. 4. Toddalia asiatica A. Habitat of liana mixed with other vegetation on medium tree estimated 12 m tall on plot 2, 1951 m, B. Colony of liana trunks, C. Habit, D. Fruit cluster HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 57 A B C D Fig. 5. Elaeagnus latifolia, A. Habitat showing liana covering estimated 10 m tall Melastomataceae tree at plot 4, 1981 m B. Liana trunk on tree, C. Habit, D. Dioecious flower 1 cm REINWARDTIA 58 [VOL.13 A C D Fig. 6. Schefflera lucida A. Habitat showing clump of secondary branches sprouded from main trunk which is climbing on large possible Lithocarpus tree, plot 5, 1981 m, B. Six or more liana trunks on large tree esti- mated 15 m tall, C. Habit, D. Flower buds B HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 59 A B Fig. 7. Vaccinium laurifolium A. Habitat of large liana on Schefflera sp. tree trunk estimated 15 m tall on plot 3, 1966 m, B. Colony of lianas at base of tree, C. Habit, D. Flower buds and fruits 1 cm 1 cm C D REINWARDTIA 60 [VOL.13 A B C D Fig. 8. Lonicera javanica A. Habitat of liana on tree trunk, B. Colony of liana trunks on plot 14, 2499 m, C. Habit, D. Single dioecious flower 3 cm HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 61 0 0.5 1 1.5 2 2.5 3 3.5 4 4.5 0 5 10 15 20 25 30 35 40 45 0 0.05 0.1 0.15 0.2 0.25 0.3 0.35 T o ta l o f l ia n a i n d iv id u a l M e a n o f li a n a d e n si ty /m 2 M e a n T ru n k d ia m e te r (c m ) 1900– 2001– 2101– 2201– 2301– 2401– 2501– Fig 9. Total of liana individual, mean of liana density/m 2 and mean trunk diameter within elevational ranges on Mt. Slamet. REINWARDTIA 62 [VOL.13 much of the work in the Pluvial Forests of the Choco Phyto–geographic Region? Such issues re- main to be answered by a Historian of Science. In the meantime, study of the high elevation lianas of Mt. Slamet, Central Java, Indonesia will continue. LITERATURE CITED AIBA S. and KITAYAMA, K. 1999. Structure, Compo- sition and Species Diversity in an Altitude Sub- strate Matrix of Rainforest Tree Communities on Mt. Kinabalu, Borneo. Plant Ecology 140: 139– 157. APPANAH, S. et. al. 1992. Liana Diversity and Species Richness of Malaysian Rainforests. J. Trop. For. Sci. 6: 116–123. ASHTON, P. F. 1964. Ecological Studies in the Mixed Dipterocarp Forests of Brunei State. Oxford For- estry Memoirs 25. ASHTON, P. F. 2003. Floristic Zonation of Tree Com- munities on Wet Tropical Mountains Revisited. Perspectives in Plant Ecology, Evolution and Sys- tematics 6: 87–104. APRIYANTI, N. 2006. Nepenthes baru dari Gunung Slamet Hingga Negeri Jiran. Trubus 435: 112–113. ARIFIANI, D. 2001. Inventarisasi Tumbuhan Ke Gunung Lawu dan Slamet, Jawa Tengah. Januari dan Oktober 2001. Laporan Perjalanan Propinsi Jawa Tengah. AVALOS, G.; MULKEY, S.S. & KITAJIMA, K. 1999. Leaf Optical Properties of Trees and Lianas in the Outer Canopy of a Tropical Dry Forest. Biotropica 31: 517–520. AVALOS, G. and AVALOS, G. 1999a. Photosynthetic Acclimation of the Liana Stigmaphyllon linde- nianum in Light Changes in a Tropical Dry Forest Canopy. Oecologia 120: 474–484. AVALOS, G. 1999b. Seasonal Changes in Liana Cover in the Upper Canopy of a Neotropical Dry Forest. Biotropica 31: 186–192. BABWETEERA, F. 2000. Effect of Gap Size and Age on Climber Abundance and Diversity in Budongo Forest Reserve, Uganda. Afr. J. Eco. 38: 230–237. BALEE, W. & CAMPBELL, D.G. 1990. Evidence for the successional status of liana forest (Xingu river Basin, Amazon Brazil). Biotropica 22: 36–-47. BEAMAN, J.H., ANDERSON, C. & BEAMAN, R.S. 2001. The Plants of Mount Kinabalu. 4. Dicoty- ledonous Families Acanthaceae to Lythraceae. Natural History Publications (Borneo) and Royal Botanic Gardens, Kew. BERLAGE, H.P. 1949. Rainfall in Indonesia: Mean Rai- fall figures for 4339 rainfall stations in Indone- sia, calculated from observations made during the period 1879–1914. De Unie. Batavia. :212. CABALLE, G. 1994. Ramet Profliferation by Longitudi- nal Splitting in the Gabonese Rainforest Liana Dalhousiea africana S.Moore (Papilonaceae). Biotropica 26: 266–275. CAMPBELL, E.J.F. and D.M. NEWBERRY. 1993. Eco- logical Relationships between Lianas and Trees in Lowland Rainforest in Sabah, East Malaysia. J. Trop. Ecol. 9: 469–490. CLINEBELL, R.; PHILIPS, O.L.; GENTRY, A.H.; STARK, N. & ZURING, H. 1995. Prediction of Neotropical Tree and Liana Species Richness from Soil and Climatic Data. Biodiversity and Conservation 4: 56–90. COLWELL, R.K.; BREHM, G.; CARELUS, C.L.; GIL- MAN, A.C. & LONGINO, J.T. 2008. Global Warming, Elevational Range Shifts and Lowland Biotic Attrition in the Wet Tropics. Science 322: 258–261. DATTARAJA, H.S. and R. SUKUMAR. 2004. Tropical Dry Forest Diversity of Mudumalai, Southern India. Inside CTFS 2004: 8–9. DAVIS, T.A. and P.W. RICHARDS. 1933–34. The Vegetation of Moraballi Creek, British Guinea; an Ecological Study of a Limited Area of Tropi- cal Rain Forest. Parts I and II. J. Ecol. 21: 350– 384; 22: 106–155. DAVIS, T.A. 1933. Biology of Plants and Animals Oc- curing in the Higher Parts of Mount Pangrango– Gedeh in West–Java. Verhandelingen der Koninklijke Akademie van Wetenschappen te Amsterdam Afdeeling Natuurkunde (Tweede Sec- tie) 31: 1–288. DILLENBURG, L.R., D.F. WHIGHAM, A.H. TERA- MURA and I.N. FORSETH. 1993. Effect of Be- low and Aboveground Competition from the Vines Lonicera japonica and Parthenosissus quinquefolia on the Growth of the Tree Host Liquidambar styraciflua. Oecologia (Berlin) 93: 48–54. GARTNER, B.L.; BULLOCK, S.H.; MOONEY, H.A.; BROWN, V.B. & Whitlock, J.L. 1990. Water Transport Properties of Vine and Tree Stems in a Tropical Decidious Forest. Amer. J. Botany 77: 742–749. GENTRY, A.H. 1982. Patterns of Neotropical Plant Spe- cies Diversity. In M.K. HECHT, B. WALLACE AND G.T. PRANCE (Eds.). Evolutionary Biol- ogy 15: 1–84. Plenum Press, New York, NY. GENTRY, A.H. 1988. Changes in Plant Community Diversity and Florsitic Composition on Environ- mental and Geographic Gradients. Ann. Miss. Bot. Gard. 75: 1–34. GENTRY, A.H. 1991. The Distribution and Evolution of Climbing Plants. In F.E. Putz and H.A. Mooney (Eds.). The Biology of Vines. :3–49. Cambridge University Press, Cambridge. GERWING, J.J.; SCHNITZER, S.A.; BURNHAM, R.J. ; BONGERS, F.; CHAVE, J.; DEWALT, S.J.; EWANGO, C.E.N.; FOSTER, R.; MARTINEZ– RAMOS, M.; PARREN, M.; PARTHASARA- THY, N.; PEREZ–SALICRUP, D.R.; PUTZ, F.E. & THOMAS, D.W. 2006. A standard Protocol for Liana Censuses. Biotropica 38: 256–261. GIODA, A., J. MALEY, R.E. GUASP and A.A. BALA- DON. 1993. Some Low Elevation Fog Forests of Dry Environments: Application to Paleoenviron- HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 63 ments. Tropical Montane Cloud Forest. Proceed- ings of an International Sysposium. HAMILTON, L.S.; JUVIK, J.O. & SCATENA, F.N. (Eds.) : 97 –101. East–West Center, Honolulu. GRUBB, P.J., LLOYD, J.R.; PENNINGTON, T.D. & WHITMORE, T.C. 1963. A Comparison of Mon- tane and Lowland Forest in Ecuador. I. The For- est Structure, Physiognomy and Floristics. J. of Ecol. 51: 567–601. GRUBB, P.J., & WHITMORE, T.C. 1966. A Compari- son of Montane and Lowland Forest in Ecuador. II. The Climate and its Effects on the Distribution and Physiognomy of the Forests. J. of Ecol. 53: 303–333. HOLDRIDGE, L. 1967. Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica. HOOVER, W. SCOTT. 1974. Begonia Collecting in Colombia. The Begonian 41 (2). HOOVER, W. SCOTT. 1980. Collecting Begonias in Papua New Guinea. The Begonian 47 (1). HOOVER, W. SCOTT. 1985. Summary of 1984 South American Collecting Expedition. The Begonian 52: 28–32. HOOVER, W. SCOTT. 1989a. Summary of 1987–88 Expedition to Ecuador. The Begonian 56: 46–50. HOOVER, W. SCOTT. 1989b. Ecuador’s Forest Ref- uge. The Explorers Journal 66: 166–169. HOOVER, W. SCOTT. 1991. Notes on a Novel Abaxial Leaf Epidermis in Ecuadorian Begonia par- viflora. J. Arnold Arboretum 71: 259–264. JACOBS, M. 1976. The Study of Lianas. Flora Malesi- ana Bulletin 29: 2610–18. KALACSKA, M., CALVO–ALVARADO, J.C. & SAN- CHEZ–AZOFEIFA, G.A. 2005. Calibration and Assessment of Seasonal Changes in Leaf Area Index of a Dry Tropical Forest in Different Stages of Succession. Tree Physiology 25: 733–744. KARTAWINATA, K. 1993. A wider view of the fire hazard. :261–266. In BROOKFIELD, H.; BYRON, Y. (Eds.). Southeast Asia’s environ- mental future: the search for sustainability. United Nations University Press, Tokyo. KARTAWINATA, K. 2005. Six Decades of Natural Vegetation Studies in Indonesia: 95–140. In SOEMODIHARDJO, S. & SASTRAPRADJA, S. D. (Eds.). Six Decades of Science and Scientists in Indonesia. Naturindo, Bogor, Indonesia. KITAYAMA, K. 1992. An Altitudinal Transect Study of the Vegetation on Mt. Kinabalu, Borneo. Vegeta- tio 102: 149–171. LONGINO, J.T. 1986. A Negative Correlation between Growth and Rainfall in a Tropical Liana. Biotropica 18: 195–200. OSAWA, M.1991.Structural Comparison of Tropical Montane Forests along Latitudinal and Altitudinal Gradients in South and East Asia.Vegetatio 97: 1 –10. OPLER, P.A., BAKER, H.G. & FRANKIE, G.W. 1994. Seasonality of Climbers: A Review and Example from Costa Rican Dry Forest. Pp. 377–391. In PUTZ, F.E. & MOONEY, H.A. (Eds.). The Biol- ogy of Vines. Cambridge University Press, Cam- bridge. PARTHASARATHNY, N.S., MUTHURAMKUMAR, S. & REDDY, M. S. 2004. Patterns of Liana Di- versity of Tropical Evergreen Forests of Peninsu- lar India. For. Eco. Manage. 190: 15–31. PEREZ-SALICRUP, D.R. 2001. Effect of liana-cutting on tree regeneration in a liana forest in Amazonia Bolivia. Ecology 82: 389–396. PHILIPS, O.L.; MARTINEZ, R.V.; ARROYO, L.; BAKER, T. R.; KILLEN, T.; LEWIS, S.L. & MALHI, Y. 2002. Increasing Dominance of Large Lianas in Amazonian Forest. Nature 418: 770– 774. PINARD, M.A. & PUTZ, F.E. 1993. Vine Infestation of Large Remnant Trees in Logged Forest in Sabah, Malaysia: Biomechanical Facilitation in Vine Suc- cession. J.Trop. For. Sci. 6: 302–309. PUTZ, F.E., LEE, H.S. & Goh, R. 1984. Effects of Post– Felling Silvicultural Treatments on Woody Vines in Sarawak. Malaysian Forester 47: 214–26. PUTZ, F.E. 1985. Woody Lianas and Forest Manage- ment in Malaysia. Commonwealth Forestry Re- view 64: 359–65. PUTZ, F.E. & CHAI, P. 1987. Ecological Studies of Li- anas in Lambir National Park, Sarawak, Malaysia. J. of Ecology 75: 523–31. PUTZ, F.E. & MOONEY, H.A. 1991. The Biology of Vines. Cambridge University Press, Cambridge. PUTZ, F.E. & WINDSOR, D.M. 1987. Liana Phenology on Barro Colorado Island, Panama. Biotropica 19: 334–341. RAHMAN, E. 2001. Eksplorasi Tumbuhan Liar Berpo- tensi obat dan Tanaman Hias de Jawa Tengah 17 –29 April. Laporan Perjalanan Propinasi Jawa Tengah. RESTOM, T.G. & NEPSTAD, D.C. 2004. Seedling Growth Dynamics of a Deeply Rooting Liana in Secondary Forest in Eastern Amazonia. For. Ecol. and Man. 190: 109–118. RICHARDS, P.W. 1936. Ecological Observations on the Rain Forest of Mt. Dulit, Sarawak. Parts I and II. J. of Ecol. 24: 1–27; 340–360. RICHARDS, P.W. 1964. The Tropical Rainforest: An Ecological Study. Cambridge University Press, Cambridge. RISWAN, S. 1982. Ecological Studies on Primary, Sec- ondary and Expermimentally Cleared Mixed Dip- terocarp Forest and Kerangas Forest in East Ka- limantan, Indonesia. Ph.D. Thesis, University of Aberdeen. RISWAN, S & KARTAWINATA, K. 1988. Regenera- tion after Distrubance in Kerangas (Heath) Forest East Kalimantan, Indonesia. :61–85. In S. SOEMODIHARDJO (Ed.). Some Ecological As- pects of Tropical Forest of East Kalimantan: A Collection of Research Reports. MAB Indonesia Contribution N. 48. RISWAN, S and KARTAWINATA, K. 1989. Regenera- tion after Disturbance in Lowland Dipterocarp Forest in East Kalimantan, Indonesia. Ekologi Indonesia 1: 1–8. RISWAN, S and KARTAWINATA, K. 1991. Species REINWARDTIA 64 [VOL.13 Guide to the Concerned Literature up to the Year 1950. Noordhoff– Kolff., Djakarta. SULISTYANI. 1995. Distribusi Paku–pakuan pada hutan lereng Selatan Gunung Slamet Baturden. KPH Banyumas Timur. Biosfera 4: 20–30. SUSANA, R. 2002. Dari kaki Gunung Slamet Menem- bus Pasar Dunia. Kompas. Selasa 9 April 2002. SVENNING, J. & CONDIT, R. 2008. Biodiversity in a Warmer World. Science 322: 206–207. TREUB, M. 1883. Observations sur les plantes Grim- pantes du Jardin Botanique de Buitenzorg. Ann. Jard. Bot. Buitenz. 3: 160–183. WEBB, L.J. 1959. A Physiognomic Classification of Australian Rainforest. J. of Ecol. 47: 551–570. WHITMORE, T.C. 1984. Tropical Rainforests of the Far East. Clarendon Press, Oxford. WHITMORE, T.C. 1998. An Introduction to Tropical Rainforests. Clarendon Press, Oxford. WIRIADINATA, H. 2004. Pengumpulan Data Tumbu- han Pengunungan de Gunung Slamet, Jawa Ten- gah 13–19 Maret 2004. Laporan Perjalanan Pro- pinsi Jawa Tengah. WRIGHT, S.J.; CALDERON, O.; HERNANDEZ, A. & PATON, S. 2004. Are Lianas Increasing in Im- portance in Tropical Forests? A 17 year Record from Panama. Ecology 85: 484–489. Strategy in Early Stage of Secondary Succession Associated with Soil Properties Status in a Low- land Mixed Dipterocarp Forest and Kerangas For- est in East Kalimantan. Tropics 1: 13–34. SADILI, A.; KARTAWINATA, K.; KARTONEGORO, A.; SOEDJITO, H. & SUMADIJAYA, A. 2009. Structure and Composition of Subalpine Summit Habitats on Mt. Gede Pangrango Complex, Cibo- das Biosphere Reserve, West Java, Indonesia. Reinwardtia 12: 391–404. SCHENCK, H. 1892–1893. Beitrage zur Biologie und Anatomie der Lianen. I. Beitrage zur Biologie der Lianen. Bot. Mitt. Trop. Hft. 4. II. Beitrage zur Anatomie de Lianen. Ibid. 5. SCHMIDT, F.H. & FERGUSON, J.H. 1951.Rainfall Types Based on Wet and Dry Period Ratios for Indonesia with Western New Guinea. Verhan- delingen Djawatan Meteorologi dan Geofisika. Djakarta 42. SCHNITZER, S.A. 2005. A Mechanistic Explanation for Global Patterns of Liana Abundance and Distribu- tion. Amer. Nat. 166: 262–276. STEENIS, C.G.G. J. VAN, HAMZAH, A. & TOHA, M. 1972. The Mountain Flora of Java. Brill, Leiden. STEENIS–KRUSEMAN, M. J. VAN. 1950. Malaysian Plant Collectors and Collections Being a Cyclope- dia of Botanical Exploration in Malaysia and a HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 65 APPENDIX 1. A. Rainfall map of section of central Java, showing Mt. Slamet, Baturaden from an old undated map providing Mean Number of Rainy days during the driest four months of the year ( Kloninklijk magnetsxh meterologisch observatorium te Batavia, gemiddeld aantal tegendagen gedurende de droogste vier maanden van het jaar). Ar- row indicating Mt. Slamet and Baturden. B. Rainfall Station Map of section of Central Java, Showing Baturaden, Mt. Slamet (Overzichtskaart van de ligging en namen der regenstation F Java & Madoera 1941). Arrow indicating station 12. REINWARDTIA 66 [VOL.13 APPENDIX 2. Circumference of Individual Lianas on Mt. Slamet Ridge# Plot #/ Species and Individual Ridge# Plot #/ Species and Individual Ridge# Plot #/ Species and Individual Ridge# Plot #/ Species and Individual Ridge 1 Plot 1 E. Pergamacea 1 2 3 4 5 6 7 8 9 10 Plot 2 T. asiatica 1 2 Plot 3 V. laurifolium 1 2 Plot 4 E. latifolia 1 2 Plot 5 S. lucida 1 Plot 6 E. latifolia 1 2 3 4 5 6 Plot 7 V. laurifolium 1 2 3 4 9 9 9.5 9 10 10 8 10.5 10 9 4.5 5 19 12 9 7.5 15.5 10 6 7.5 10 9 8 10 10.5 9 8 5 6 7 E. latifolia 1 2 Plot 8 V. Laurifolium 1 S. lucida 1 T. Asiatica 1 Plot 9 T. Asiatica 1 V. laurifolium 1 2 3 4 5 6 7 Plot 10 V. laurifolium 1 2 3 4 5 6 7 8 S. lucida 1 2 3 4 5 6 7 8 9 10 7.5 5 6 4.5 6 14.5 15 10 14 6.5 6 10 6.5 11 7 6 7 7.5 8 8.5 7 7.2 7.2 8 5 6 6 6.5 7 6.5 8 9 11 12 Plot 11 V. laurifolium 1 2 3 T. asiatica 1 2 3 4 5 Plot 12 V. laurifolium 1 2 3 4 5 6 Plot 13 V. laurifolium 1 2 3 4 5 6 7 8 9 Plot 14 L. javanica 1 2 3 4 5 6 7 8 9 10 13.5 17.5 5.5 10.5 7 5.5 5 5 20 12 8.5 7.5 18.5 9.5 8 7 6.5 21 15.5 5.5 5.5 6 9.5 7 5.5 8 4.5 10 6 9.5 8 9.5 6 Plot 15 L. javanica 1 2 3 Plot 16 V. laurifolium 1 2 Plot 17 V. laurifolium 1 2 3 4 5 6 7 Plot 18 V. laurifolium 1 2 3 4 5 L. javanica 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 8.5 10 8.5 17 16.5 10.5 5.5 12 15.5 7.5 7.5 18.5 24 10 12.5 16.5 5.5 11.5 8 7 8.5 5 5 5.5 4.5 5 4.5 5.5 5 4 10.5 4.5 7.5 6 5.5 7 7 6.5 8 C ir cu m fe re n ce ( cm ) C ir cu m fe re n ce ( cm ) C ir cu m fe re n ce ( cm ) C ir cu m fe re n ce ( cm ) HOOVER : Exploration of high elevation liana colonies on Mt. Slamet, Central Java, Indonesia 2009] 67 Ridge# Plot #/ Species and Individual Ridge# Plot #/ Species and Individual Ridge# Plot #/ Species and Individual Plot 19 V. laurifolium 1 2 3 4 5 6 7 15.5 12 13 12 6.5 8.5 6 Plot 20 V. laurifolium 1 Plot 21 V. laurifolium 1 18 64 Plot 22 L. javanica 1 2 3 4 5 6 6.5 6.5 6 6 6 6.5 C ir cu m fe re n ce ( cm ) C ir cu m fe re n ce ( cm ) C ir cu m fe re n ce ( cm )