Vascular plant species inventory of a Philippine lowland rain forest ...

Biodiversity and Conservation (2006) 15:1271–1301 Ó Springer 2006DOI 10.1007/s10531-005-2576-4-1Vascular plant species inventory of a Philippinelowland rain forest and its conservation valueGERHARD LANGENBERGER, KONRAD MARTIN * andJOACHIM SAUERBORNInstitute of Plant Production and Agroecology in the Tropics and Subtropics (380), AgroecologySection, University of Hohenheim, 70593 Stuttgart, Germany; *Author for correspondence (e-mail:martin-k@uni-hohenheim.de; phone: +49-711-459-3605; fax: +49-711-459-3843)Received 22 June 2004; accepted in revised form 8 February 2005Key words: Conservation value, Dipterocarp forests, Gene bank, Molave forest, Native species,Species richness, Tropical rain forest, Vascular plant speciesAbstract. The Philippines are one of the most important biodiveristy hotspots on earth. Due to theextraordinary rate of environmental destruction, leaving only 3% of the land with primary forest,this biodiversity is at high risk. Despite that situation information on Philippine forest vegetation isfragmentary and focused on trees. This study aimed at analysing forest remnants in the LeyteCordillera on the Island of Leyte, and at evaluating their role as refuge to the largely destroyedlowland forest vegetation. A total of 49 plots (100 m 2 each) between 55 and 520 m a.s.l. werestudied. All vascular plant species except epiphytes were included. Records include 685 taxa from289 genera and 111 families, representing nearly 8% of the known Philippine vascular plant species.More than half (52%) of the species are Philippine endemics. A number of 41 tree species, or 6% ofall taxa recorded, are included in the IUCN red list, either as vulnerable, endangered, or criticallyendangered. Life form composition was dominated by phanerophytes (65.3%), followed by lianasand chamaephytes (17.1 and 16.9%, respectively). The most common families were the Rubiaceaewith 35 and the Euphorbiaceae with 32 species. All five Philippine dipterocarp forest types as wellas the molave forest type were represented by typical tree species. The area provides an importantgene bank of the highly threatened Philippine lowland forest vegetation and is of high value forbiodiversity conservation. Additionally, it can play an important role as seed source of valuable treespecies for the increasing initiatives to rehabilitate and reforest degraded land with native species.IntroductionThe destruction of tropical rain forests is still continuing at high rates (FAO2003). This process, especially threatens the earth’s biodiversity hotspots such asthe Philippines (Myers et al. 2000; Brooks et al. 2002). Despite this, there are onlyvery few studies worldwide which aimed at the documentation of the total plantspecies richness of such sites. Most inventories were restriced to selected life formssuch as ground herbs (e.g. Kiew 1987; Poulsen and Balslev 1991; Poulsen 1996) ortrees of a defined minimum diameter (e.g. Valencia et al. 1994; Lieberman et al.1996; Newbery et al. 1996; Rennolls and Laumonier 2000; Slik et al. 2003).Vascular plant species composition of tropical lowland forests was studied inGhana on 0.5- and 1-ha plots by Hall and Swaine (1981), in Amazonia on 0.02-ha plotsby Takeuchi (1960) and on 10 non-contiguous plots of 0.1 ha by Duivenvoorden

1272(1994), in Ecuador on 0.1-ha plots of three lowland forest types by Gentry andDodson (1987), and in stratified plots with a total area of about 2 ha in PuertoRico by Smith (1970). In Southeast Asia, Kochummen et al. (1992) studied thetrees and shrubs (>1 cm diameter at breast height (dbh)) in a 50-ha plot in thePasoh Forest Reserve in Malaysia. The most comprehensive study including allvascular plants as well as mosses was conducted by Whitmore et al. (1985) on asingle 100-m 2 plot in the lowland rain forest of Costa Rica. However, no studyrepresenting a complete inventory of vascular plant species richness of any site oflowland rain forest in Southeast Asia was found.The Philippines are among the most seriously depleted tropical countrieswith only 3% of the land area still covered by primary forest (Myers et al.2000). From 1990 to 2000, the Philippines lost 1.4%, or 89,000 ha, of the forestarea annually (FAO 2003). At the same time, the Philippine archipelago is oneof the most important biodiversity hotspots on earth (Myers et al. 2000) withhigh proportions of endemic plant and animal species (Heaney and Regalado1998). The endemism rate of plants was estimated to be 39% (Davis et al.1995), but for certain taxa, it can be much higher. For example, 11 of 12 speciesof pitcher plants (Nepenthes spp.) known from the Philippines are endemic(Cheek and Jebb 2001). Similarly, there are high rates of endemism among thefauna. Referring to terrestrial vertebrates, 64% of the archipelago’s landmammals are endemic, as well as 44% of the breeding land birds, 68% of thereptiles, and ca. 78% of the amphibians (Heaney and Regalado 1998). Most ofthem depend on forest ecosystems.Despite the ecological uniqueness on the one hand and the extensivedestruction on the other, the study of Philippine forest vegetation has beenneglected (Tan and Rojo 1989; Kartawinata 1990; Soerianegara and Lemmens1994). Much of the current knowledge is still based on studies conducted in theearly 20th century (Whitford 1906; Whitford 1911; Brown and Mathews 1914;Brown 1919), which were mainly dealing with timber trees under economicalaspects. Recent studies focused on the vegetation of montane and submontaneforest types on different islands. However, in most cases (Aragones 1991;Pipoly and Madulid 1998; Proctor et al. 1998; Hamann et al. 1999) these werelargely restricted to trees of a defined size, which usually is ‡10 cm dbh. Buotand Okitsu (1997) only considered woody plants higher than 1.3 m, and Ingle(2003) those of at least 5 cm dbh. The only data without size limitations areprovided by Gonzales-Salcedo (2001) from Mt. Amuyao, Luzon, at elevationsof 1600–1800 a.s.l. and by Gruezo (1998) from the highly degraded vegetationof Pagbilao Grande Island. No study dealing with lowland forest vegetationwas found in the literature.In order to provide more substantial information on species richness andcomposition of Philippine lowland forests, we analysed forest remnants in therugged foothills of the Leyte Cordillera. The island of Leyte is located in thecentral part of the Philippine archipelago and represents a typical example ofthe environmental situation in the Philippines. In 1987, the remaining forestcover of Leyte was 12%, and in 1994 only 2% of the island’s area have been

estimated to be primary forest (Dargantes and Koch 1994). More recent data(DENR 1998) show that about 40% of the land area of Leyte is covered bygrassland and barren land, resulting from abandoned cultivation and grazingland that marginalised in productivity through erosion and leaching. Another40% of the island’s area is under coconut plantations. The remaining area iscomposed of settlements, agricultural land and forest. In the view of this situation,the objectives of this study were (a) to analyse the vascular plant speciescomposition and diversity of selected plots of mature primary forest and (b) toevaluate the role of the study area as refuge to lowland forest vegetation and itssignificance for conservation and as a gene bank.Material and methodsStudy area1273The Island of Leyte (Figure 1) belongs to the biogeographic region of theEastern Visayas (DENR and UNEP 1997). It is located between 9°55¢ N–11°48¢ N and 124°17¢ E–125°18¢ E, with an extension of 214 km from north tosouth. Located offshore the northeastern part of Leyte is the island of Samar.The southern part of Leyte is exposed to the Pacific Ocean (Leyte Gulf).Leyte is characterised by the north–south running Leyte Cordillera which ispart of the Philippine Fault Line. The Cordillera reaches a maximum elevation ofca. 1350 m (Mt. Lobi) in the northern part of the island. As geologically youngvolcanic mountain range, it shows a typical rugged topography of narrow ridgesand steep slopes, where landslides are common (Bremer 1995, 1999; Walsh 1996).In its foothills, patches of primary forest without discernible human interferencecan still be found although the island is densely populated (ca. 262 inhabitants/km 2 as calculated after: NSCB RU-8 2001). The coastal plains have already beendeforested in the first half of the last century (Barrera et al. 1954).The study area is located in the western part of the Leyte Cordillera,ca. 8 km north of the provincial capital of Baybay, in the foothills of Mt.Pangasugan (1150 m, 10°46¢ N, 124°50¢ E). In this part of the island, theCordillera reaches close to the coast. Large parts of the mountain’s westernrange are extremely steep and are free from trees of this reason. In the easternpart of the Cordillera, the slope has a lower gradient.Primary forest can be found from about 250 m a.s.l. up to the mountain’ssummit at 1150 m a.s.l. In hillsides below this elevation, the forests has largelybeen replaced by coconut plantations and slash and burn agriculture. Onlyalong the small creeks, near natural vegetation can still be found at these lowerelevations. Above ca. 600 m a.s.l., the lowland forest formation changes intomossy forest, with its stunted trees and a rich epiphyte community.Within the study area, no recent logging was observed, although forestclearing continues at other localities of the mountain. This can be explained bythe area’s status as Forest Reserve of the Leyte State University. Despite this,rattan collection and hunting was observed within the Forest Reserve.

1274Figure 1.The Philippine archipelago, the Island of Leyte, and the location of the study area.Geology and soilsThe soil type in the primary forest between 370 and 520 m a.s.l. is a haplicAndosol with rudic phase (FAO/UNESCO 1988) overlying basaltic andandesitic breccia (Zikeli 1998). The soil at lower elevations (100 m a.s.l.) hasbeen classified as haplic Alisol (FAO/UNESCO 1988) over basalt (Asio 1996).ClimateAccording to the climatic classification of the ‘Modified Corona’s System’,Leyte is climatically divided (Kintanar 1984). Southern Leyte belongs to theclimatic type II (i.e. no pronounced dry season), but exhibits a distinct rainfallpeak in December and January as a result of the northeast monsoon. Thenorthern part of Leyte which includes the study area has been assigned toclimatic type IV, showing a more or less even rainfall distribution throughout

1275the year. The standardization of the rainfall pattern in northern Leyte comparedto that of southern Leyte might be explained by the protective effect ofthe Island of Samar off the northeast-coast of Leyte, although Samar’smountains are lower than those of Leyte (ca. 850 m a.s.l.).Local climatic conditions have been analysed from a 23-years period (1976–1998) of record by using data from the PAGASA (Philippine Atmospheric,Geophysical and Astronomical Services Administration) weather station onthe campus of the Leyte State University (7 m a.s.l.), ca. 1–3 km west of thestudy plots. The annual average temperature is 27.4 °C and the average annualprecipitation is 2586 mm. Highest precipitation occurs during November toJanuary. Lowest rainfall is observed between March and May. On average, allmonths receive at least 100 mm precipitation, i.e. there is no dry monthaccording to the definition of Walsh (1996). However, drought periods (i.e. lessthan 50 mm of monthly precipitation according to Walsh 1996) of up to4 months have been recorded during El Nin˜ o Southern Oscillation events. Thegeneral rainfall patterns and the climatic conditions measured at the PAGASAstation are more similar to climatic type II with its clear impact of the northeastmonsoon than to climatic type IV, implying that neither the mountainsof Samar nor the Leyte Cordillera itself causes a distinct rain shadow westof Mt. Pangasugan.Orographic rainfalls are an important factor in the Leyte Cordillera, especiallyin the vicinity of the mountain summits. The summit of Mt. Pangasuganis often observed being cloud covered, and during field work heavy rainfallshave been experienced, while the coastal plain did not receive any precipitation.An important climatic feature of the area are typhoons. Leyte lies at thesouthern margin of the typhoon tracks entering the Philippines, and is hit at arate of five typhoons in three years, mainly during the summer months (Parong1984; cited in Kintanar 1984).Vegetation analysisField studies were conducted in 1997 and 1998. The attempt to identify aminimum area in mature primary forest failed due to the heterogeneity of thevegetation. A plot size allowing a reasonable number of replications proved tobe 100 m 2 . Where relief conditions and homogeneity of the vegetation allowed,plots were arranged along a catena from ridge to river bank. The 100-m 2 plotswere generally designed as quadrats, but on narrow ridges and river banks,other rectangular design was used due to relief constraints. A total of 49 plotswas established, with 15 on the ridge, 21 on slopes, and 11 on riverbanks. Twoplots were established in ca. 6-year-old land slide successions, with one of themlocated ca. 2 km south of the main study area.The vegetation analysis procedure was based on a ‘nested quadratdesign’ (Kent and Coker 1992). All plants >2.5 m were identified fromthe total plots (100 m 2 ). On central subplots of 25 m 2 , all plants £ 2.5 m

1276as well as the lianas were considered. Records included epiphytic and climbingplants on the stem bases of trees up to a height of 2.5 m. Crown epiphytes werenot included in the analysis, but epiphytes found on the ground were identifiedand added to the species lists. From species which could not be identified in thefield, voucher specimens were collected. Tall trees were sampled with the helpof a tree climber. However, no samples were taken from erect and climbingpalms (rattans), because this would have been destructive and the chance ofidentification was very low due to the lack of fertile specimens. Therefore, mostpalms had to be distinguished as morphospecies.Taxa were assigned to life forms on the base of field observations, or with thehelp of literature information in the case of juveniles. Life form classificationfollowed Ellenberg and Mueller-Dombois (1967). Plant samples collected inthis study were deposited at the Department of Plant Breeding, Herbarium,Leyte State University, ViSCA, Baybay, Leyte, 6521A, Philippines.Species identification and nomenclatureIdentification of specimens was conducted with the help of literature andspecialists. Publications referring to the Philippine flora included de Guzmanet al. (1986), Merrill (1912), Pancho (1983), Santos et al. (1986), van Steenis(1950-ongoing), and Zamora and Co 1986). Sources referring to neighbouringcountries, which also include many Philippine taxa, were used in addition(Henderson 1974a, b; Ng 1978, 1989; Keng 1983, 1990; Whitmore 1983a, b;Corner 1988; Anonymous 1993, 1994, 1996; Soepadmo and Wong 1995;Soepadmo et al. 1996). Identification of seedlings, infertile and juvenile plantswas not possible in all cases. For pre-identification of taxa and delimitation ofmorphospecies, field characters were very important. Besides the description ofsuch characters in the above mentioned literature, the specific publications byvan Balgooy (1997a, b) and Keller (1996) were used. Tree seedlings wereidentified with help of Ng (1991) and Burger (1972).In addition, plants were identified by taxonomists from the PhilippineNational Herbarium, Manila, the National Herbarium of the Netherlands,Leiden Branch, and collaboratively during meetings of the Philippine NativePlant Group. Nomenclature followed various sources as cited above. However,priority was given to the Flora Malesiana (van Steenis 1950-ongoing), wheneverpossible. The legumes were assigned to the traditional family ofLeguminosae. Scientific names were not derived from the translation of localnames in any case.Data analysisSpecies richness, diversity and evenness were determined for each of the 49plots. Only those plants rooting within the plots were considered in the

1277analysis. The Shannon-Index (H¢) was used as a robust and simple diversitymeasure (Magurran 1988). For the analysis of species dominance patterns,Evenness (E) based on the Shannon-Index was calculated for each of the plots.To assess the area’s value as a refuge to Philippine tree species, the characterizationof forest types by Whitford (1911) was used. His classification andcharacterization is based on the occurrence of typical tree species and treespecies combinations. He often used vernacular names or typical families orgenera as e.g. ‘Apitong’ for Dipterocarpus spp. to characterise his forest types.For many of these vernacular names a scientific species could not be assignedwith certainty, and therefore, were not used for comparisons. Whitford (1911)pointed out that the description of his forest types was based on a still fragmentaryknowledge of Philippine forests. Most of his ‘typical’ tree species –with few exceptions such as mangroves – occur in the other forest types as well.For example, many species of the dipterocarp forest types occur at wet localitiesin the Molave forest (limestone forest). On the other hand, the typicalMolave forest species also exist in the dipterocarp forest types, especially ondry sites. Of such reasons, Whitford’s (1911) forest types are primarily relatedto the major habitat conditions in the Philippines and do not represent realplant associations. The comparison of the species recorded in this study withWhitford’s (1911) forest types merely demonstrates the diverse habitat conditionsin the present study area. Unfortunately, not much work has been conductedso far to improve Whitford’s system, and information on speciescomposition of the undergrowth vegetation, which might be especially valuableto characterise habitat conditions (Schulze and Whitacre 1999), is still missing.ResultsFrom the 49 plots, a total of 685 taxa was recorded. Of these, 58.3% wereidentified to species level, 86.2% to genus level, and 96.7% to family level. Theremaining 3.3% of the taxa could only be assigned to higher taxonomic levels.All taxa identified to species level are listed in the Appendix. Species inventorywas clearly dominated by angiosperms, accounting for 92.1% of all species.The pteridophytes represented 7.5% of the species. Only three species ofgymnosperms (Podocarpus rumphii, Gnetum gnemon, G. latifolium) were found(Table 1).More than half (52%) of all species identified are endemic to the Philippines,including one endemic genus (Greeniopsis, Rubiaceae). The most commonfamilies were the Rubiaceae (35 species) and the Euphorbiaceae (32 species),followed by the herbaceous family of Araceae and the erect and climbing palms(Arecaceae) with 28 species each. The Meliaceae and Moraceae included 27species each (Figure 2). The ratio between the number of genera and thenumber of species ranged between 1:1.5 (Anacardiaceae) and 1:6.7 (Moraceae).The frequency of taxa was low. Nearly half (48.5%) of all taxa were recordedfrom only one of the 49 plots, and nearly one third (30.5%) of the taxa were

1278Table 1. Taxonomic composition of 49 non-contiguous plots (100 m 2 each) in lowland forestremnants of the study area in the foothills of Leyte Cordilliera.Spermatophytes (%) Pteridophytes (%) Total (%)GymnospermsAngiospermsFamilies 2 (1.8) 94 (84.7) 15 (13.5) 111 (100)Genera 2 (0.7) 261 (90.3) 26 (9) 289 (100)Species 3 (0.4) 631 (92.1) 51 (7.5) 685 (100)Figure 2. The 20 most common plant families recorded from 49 plots (100 m 2 each) in the studyarea. Figures in brackets indicate the ratio between the number of genera and the number ofspecies.

1279represented by only one single individual. Very few species showed high frequenciesas e.g. the two tree species, Calophyllum blancoi (present in 32 plots)and Dacryodes rostrata (present in 31 plots), which was due to a high rate ofjuveniles.The average number of species per plot was 47 and ranged between 17 and80. Shannon diversity (H¢) reached values between 2.2 and 3.9, and evenness(E) ranged between 0.64 and 0.98. The species–area curve for all plots shows asteady increase of species numbers with only a weak tendency to level off(Figure 3). The flattening of the curve at its beginning is the result of the riverbank vegetation which was comparatively species poor and homogenous. Thespecies–area curve starts to rise again with the addition of the slope plots.Life form composition is clearly dominated by phanerophytes (65.3% of alltaxa), followed by lianas (17.1%) and chamaephytes (16.9%). Geophytes wererare (0.7%) and largely represented by few ground orchids. Hemicryptophytesand therophytes were absent (Figure 4). Epiphytes were not the focus of thisstudy and are therefore not included in the calculation of life form composition.A rough estimate of epiphyte contribution to the area’s species inventoryis ca. 10%. The most conspicuous epiphytic plant group observed were orchids.Many of the vegetation clusters observed in the tree crowns were composedof the accumulation of orchid bulbs belonging to a single species (e.g.Grammatophyllum multiflorum).The following taxa occurring in the study plots have been classified bySoepadmo (1995) as endangered and economically important lowland forestgenera in SE Asia: Anisoptera, Dipterocarpus, Parashorea, Shorea, Vatica(Dipterocarpaceae), Artocarpus (Moraceae), Mangifera (Anacardiaceae), andCalamus (Arecaceae). Additionally, 41 of the tree species recorded are listed asendangered for the Philippines by IUCN (2000). Of these, 23 are classified asvulnerable, one as endangered, and 17 as critically endangered (see Appendix).Figure 3.Species–area curve for 49 plots (100 m 2 each) in the study area.

1280Figure 4. Life form spectrum (after Ellenberg and Mu¨ller-Dombois 1967) of species recordedfrom 49 plots (100 m 2 each) in the study area.DiscussionThe 685 taxa recorded from the 49 plots account for nearly 8% of the ca. 8900vascular plant species so far described for the Philippines (Davis et al. 1995).Although the plots were not contiguous and species numbers can therefore beexpected to be higher than in contiguous plots (Whitmore 1985) this figure ishigh, considering the small overall study area (4900 m 2 in total). Only very fewdatasets cover tropical lowland forest vegetation comprehensively and aretherefore suitable for comparison. The only study using the same plot size wasconducted by Whitmore et al. (1985) in the tropical lowland rain forest ofCosta Rica, who analysed a plot of 100 m 2 , considering all vascular plants.They recorded a total of 233 species, including 59 (25%) epiphyte species. Inthe present study, the highest number of species recorded from a single 100-m 2plot was 80 and thus much lower than the number found by Whitmore et al.(1985). However, in the present study the vegetation up to 2.5 m tall as well asthe lianas were collected from subplots of 25 m 2 , and crown epiphytes wereexcluded. Despite this, the maximum number of vascular plant species on 100-m 2 plots in the study area can expected to be clearly lower than the number of233 species recorded by Whitmore et al. (1985).An estimate of the overall vascular plant species richness of Mt. Pangasuganarea, including mossy forest as well as the different stages of succession, resultsin 1500–2000 species. This estimate is based on the very conservativeassumption that 50% of the lowland forest species was recorded in this study,and that the mossy forest has a similar species richness as a 1-ha plot studied by

1281Meijer (1959) in a montane rainforest in Indonesia (333 vascular plant species).The numbers of tree species given by Ingle (2001) (100 species ‡5 cm dbh) andHamann et al. (1999) (92 species ‡10 cm dbh) for 0.75- and 1-ha plots,respectively, in Philippine mountain environments show that the overall speciesrichness including all life forms can be expected to be roughly similar to that ofMeijer (1959) in Indonesia.Representation of taxaThe composition of taxa observed in this study is similar to other areas inSoutheast Asia. Differences to such sites are related to the proportions offamilies and result mainly from different inventory approaches. For treesalone, the dominance of the Dipterocarpaceae and the Euphorbiaceae concerningnumber of species is well documented (Manokaran and Kochummen1990; Sist and Saridan 1998; Slik et al. 2003; Wilkie et al. 2004). Sist andSaridan (1998) report that the Dipterocarpaceae represent 70% of alltrees ‡ 50 cm dbh in a primary forest in East Kalimantan. In our study, theDipterocarpaceae were the most common family in the canopy layer (12 of 44species).Turner (1994) analysed the vascular flora of Singapore and its main habitattypes from herbarium collections.The Orchidaceae are the most speciose familyin his taxonomic spectrum. This reflects the large number of orchid species inMalesia (6500 species according to Soepadmo 1995). In our study, however,Orchidaceae are poorly represented because we did not include crown epiphytes.Without considering the orchids in both studies, the pteridophytes arethe most speciose group, followed by the Rubiaceae and the Euphorbiaceae inboth studies. The other predominant families in terms of species richness listedby Turner (1994) are Annonaceae, Moraceae, Arecaceae/Palmae, Myrtaceae,Melastomataceae and Lauraceae. With exception of the Melastomataceae,these are also the most speciose families in our study (Figure 2).Representation of forest typesThe rugged relief of the study area represents a broad spectrum of Philippinehabitats. The comparison of the tree species recorded from our study with thetypical tree species composition of the forest types described by Whitford(1911) showed a high degree of correspondence. Many tree species typical ofthe five dipterocarp forest types as well as the Molave type (Figure 5) werepresent. From the 18 tree species listed by Whitford (1911) as typical for theLaua´n-haga´ghak, 15 were also present in our study area. Originally, this foresttype is established on lowland plains on wet soils (Whitford 1911), but wastransformed into rice fields in the study area. However, tree species representingthis type of forest still occur on the banks of the small creeks at low

1282Figure 5. Comparison of the number of characteristic species of the different lowland forest typesin the Philippines (after Whitford 1911) with the number of respective species recorded in thisstudy.elevations. The typical tree species of other forest types were also well represented(Figure 5). The high number of Molave type species (50% of the typicalspecies as mentioned by Whitford 1911) in the study area is remarkable, as thisforest represents dry limestone areas (Whitford 1911). This is another indicationthat the area’s vegetation might be strongly influenced by drought periods.Life form compositionThe dominance of trees and phanerophytes is a typical feature of tropical rainforests (Richards 1996). In our study, Phanerophytes account for 65.3% of thespecies. Richards (1996) provides figures from a rain forest in Guyana, whichare based on the Raunkiaer System (Raunkiaer 1934) and cannot be directlycompared with our data. We therefore recalculated his data by excluding thelianas from the phanerophytes and excluded the epiphytes in addition. Thisresulted in a life form composition of 60% phanerophytes, 16% chamaephytes,and 24% lianas. A similar recalculation of figures provided by Cromer andPryor (1942) for a rain forest in Queensland results in 77.1% phanerophytes,12.5% chamaephytes, and 10.4% lianas. Figures for a terra firme rain forest in

1283Brasilia (Cain et al. 1956) are: phanerophytes 74.3%, chamaephytes 0.9%,hemicryptophytes 2.8%, geophytes 0.9%, lianas 12.8%, and epiphytes 8.3%.Therophytes and hemicryptophytes are usually absent from undisturbedtropical rain forests (Richards 1996). Geophytes are also often absent as inRichards’ Guyana study or represented by few species as in the present study(0.7%), where they were mainly made up of ground orchids.The estimated proportion of epiphyte species of the total number of speciesin this study (ca. 10%) is clearly lower than the numbers given by Whitmoreet al. (1985) (25%) for Costa Rica or by Gentry and Dodson (1987) (35%) forEcuador.Conservation valueKochummen et al. (1992) stated that comparatively small areas might representhigh numbers of a regional flora. They found that their 50-ha plot in the PasohForest Reserve (Malaysia) included 25% of all trees and shrubs (‡ 1 cm dbh)of the Malay Peninsula. In our study, an overall sample area of approximatelyhalf hectare included ca. 8% of all Philippine vascular plant species. Given thesmall area considered as well as the fact that neither the successional vegetationnor the mossy forest is included, the representation of Philippine flora in theMt. Pangasugan area is clearly higher than 8%.The proportion of 52% endemic taxa recorded in this study is clearly higherthan the proportion of 39% stated as an average for the Philippines (Daviset al. 1995). This result agrees with Ashton (1993) who stated that the southeasternpart of the Philippines is especially rich in endemic plants. The area’sendemism might be even higher than 52%, as a number of taxa could not beidentified. For example, only 3 of the 16 rattan species (Arecaceae) recorded,which generally show a high degree of endemism (Dransfield 1990), could beassigned to a scientific name. Two of them were Philippine endemics.Another aspect referring to the conservation value of the area is theoccurrence of 41 tree species in the red list of IUCN (2000). However, from thespecies recorded from this study, other than trees are not represented in the redlist. Despite this, it can be expected that many of the non-tree taxa recorded arethreatened by habitat destruction as well. For example, no rattans are listed byIUCN although this plant group is still heavily exploited and shows high ratesof endemism. The IUCN red list seems to have a strong focus on well knownand economically important tree species. This is supported by the fact that onlydipterocarps are classified as critically endangered, although many other treespecies are more rare in the study area. This was e.g. true for the valuable treespecies Heritiera sylvatica (Sterculiaceae) and Xanthostemon verdugonianus(Myrtaceae) which were known by local farmers from only one mature treeeach in the entire western foothills of Mt. Pangasugan.Taken together, the Mt. Pangasugan region on Leyte represents a uniquerefuge for a high number of species, which are characteristic of all Philippine

1284dipterocarp forest types and the molave type. In view of the large areas ofdegraded land in the Philippines, the conservation value of the Mt. Pangasuganregion is very high and represents an important gene bank of the Philippineforest vegetation.AcknowledgementsThis study was part of the ViSCA-gtz Applied Tropical Ecology Program (PN95.2290.5–001.00) and partly funded by the Tropical Ecology Support Program(TO¨ B) of GTZ (PN 90.2136.1–03.107). We are grateful to the Presidentof the Leyte State University, Dr. P.P. Milan and her staff for their support andhelp. We are also grateful to the Cienda San Vicente Farmer Association(CSVFA) and their community organiser Marlito Bande, who made theextensive field trips possible. Special thanks to the curator of the PhilippineNational Herbarium, Dr. Madulid, and his staff, as well as to the director ofthe National Herbarium of the Netherlands, Leiden branch, Prof. Baas and hisstaff who helped to put species identification on a firm ground. We are alsovery grateful to Leonardo Co, Nina Ingle, David and Luze Bicknell, FranzSeidenschwarz, and B.C. Tan for various assistance. We also like to thank thetwo anonymous referees for their comments and constructive criticism.AppendixSpecies list of the vascular plant species found in 49 plots (100 m 2 each) in thefoothills of the Leyte Cordillera at Mt. Pangasugan, Leyte, Philippines.The list includes only those species which could be identified to species level.Some species recorded outside the plots are provided in addition.Numbers in brackets following the species name indicate the first voucherspecimen collected of this species.Life form classification of species is based on observations of mature individualsin the study area, or from species descriptions in literature. Life formdefinitions follow Ellenberg and Mueller-Dombois (1967) with a minor revisionby Richter (1997). MacP, Macrophanerophyte (>20–50 m); MesP, Mesophanerophyte(>5–20 m); MiP, Microphanerophyte (>2–5 m); NP, Nanophanerophyte(>1–2 m); NP herb, herbaceous Nanophanerophyte; Ch,Chamaephyte ( £ 1 m); Ch frut, fruticose Chamaephyte; Ch suff, suffruticoseChamaephyte; Ch herb, herbaceous Chamaephyte; G rhiz, rhizome Geophyte;PL, Phanerophytic Liana; r PL, root PL; st PL, winding PL; el PL, tendril PL; dPL, spread climber; E, Epiphyte.

Species classified by IUCN (2000) as endangered are listed along with theirstatus in bold letters. Short definitions of the status are:CR, critically endangered (‘… facing an extremely high risk of extinction inthe wild in the immediate future …’); EN, endangered (‘… not criticallyendangered but facing a very high risk of extinction in the wild in the nearfuture …’); VU, vulnerable (‘… not critically endangered or endangered butfacing high risk of extinction in the wild in the medium-term future …’).For comprehensive definitions and criteria of classification see www.iucnredlist.org/search-basic.htmlI.1285SpermatophytaAceraceaeAcer laurinum Hassk. (1221)ActinidiaceaeSaurauia cf. denticulata C.B. Rob. (1078)Saurauia samarensis Merr. (235)AlangiaceaeAlangium longiflorum Merr. (1331) VUAmaranthaceaeDeeringia polysperma (Roxb.) Moq. (2214)AnacardiaceaeDracontomelon dao (Blco.) Merr. & Rolfe (660)Dracontomelon edule (Blco.) SkeelsKoordersiodendron pinnatum (Blco) Merr. (162)Mangifera altissima Blco. (971) VURhus taitensis Guill. (818)Semecarpus cuneiformis Blco. (538)AnnonaceaeAlphonsea arborea (Blco.) Merr. (1009)Anaxagorea javanica Bl. (1509)Artabotrys cf. rolfei Vid. (2159)Cananga odorata (Lamk.) Hook. f. & Thoms.Goniothalamus elmeri Merr. (327)Meiogyne virgata (Bl.) Miq.Papualthia cf. lanceolata (Vid.) Merr. (206)Popowia pisocarpa (Bl.) Endl. (1054)ApocynaceaeAlstonia macrophylla Wall. ex. G. Don (1774)Alstonia scholaris (L.) R. Br.Kibatalia blancoi (Rolfe) Merr. (467)Lepiniopsis ternatensis Val. (2220)Tabernaemontana pandacaqui Poir. (140)Voacanga globosa (Blco.) Merr. (218)AraceaeAlocasia cf. zebrina Schott ex van Houtte (1677)Amorphophallus paeoniifolius (Dennst.) Nicolson (1924)Costus speciosus (J. Konig) SmLife formMacPMiPMiPMesPCh herbMacPMesMacPMacPMesPMiPMesPMiPel PLMesPMiPMesPMesPMesPMesPMesPMesPMesPMicPMiPCh herbNP herbNP herb

1286SpermatophytaPothos cylindricus Presl (1226)Raphidophora korthalsii Schott (881)AraliaceaeArthrophyllum ahernianum Merr. (1911)Osmoxylon trilobatum (Merr.) Philipson (220)Polyscias nodosa (Bl.) Seem.ArecaceaeCalamus cf. merrillii Becc.Caryota cf. cumingii Lodd. ex MartCaryota cf. mitis Lour. (560)Daemonorops cf. mollis (Blco.) Merr. (593)Korthalsia laciniosa Mart. (1120)Pinanga maculata PorteCaryota rumphiana Mart. var. philippinensis Becc.AristolochiaceaeAristolochia philippinensis Warb. (702)AsclepiadaceaeHoya multiflora Bl. (689)AsteraceaeVernonia arborea Buch.-Ham. (826)BignoniaceaeOroxylum indicum (L.) Vent.Radermachera pinnata (Blco.) Seem. (1129)BurseraceaeCanarium asperum Benth. (265)Canarium denticulatum Bl. (428)Canarium euryphyllum Perk. (1265)Canarium gracile Engl. (611)Canarium hirsutum Willd. (1714)Dacryodes rostrata (Bl.) H. J. Lam (247)CaprifoliaceaeSambucus javanica Reinw. ex Bl.CasuarinaceaeGymnostoma rumphianum (Miq.) L.A.S. Johnson (1915)CecropiaceaePoikilospermum erectum (Blco) Merr. (321)Poikilospermum suaveolens (Bl.) Merr. (328)CelastraceaeBhesa paniculata Arn. (812)Euonymus cochinchinensis Pierre (495)Euonymus javanicus Bl. (232)Lophopetalum javanicum (Zoll.) Turcz. (1127)ChloranthaceaeChloranthus erectus (Buch.-Ham.) Verdc. (1522)Sarcandra glabra (Thunb.) Nakai (562)Life formr PLd PLMesPNPMesPd PLMiPMesPPLd PLMiPMacPCh suffPLMesPMesPMesPMacPMesPMacPMesPMesPMacPMiPMesPd PLd PLMesPMesPMesPMacPCh suffCh frut

1287SpermatophytaChrysobalanaceaeMaranthes corymbosa Bl. (790)ClusiaceaeCalophyllum blancoi Pl. & Tr. (278)Calophyllum soulattri Burm. f. (1250)Cratoxylum formosum Benth. & Hook. f. ex Dyer (452)CombretaceaeTerminalia microcarpa Decne. (672)Terminalia nitens Presl (481) VUCommelinaceaeFloscope scandens Lour.Forrestia hispida Less. & A. Rich. (423)Pollia sorzogoniensis (E. Meyer) Steud.Pollia thyrsiflora (Bl.) Steud.Rhopalephora cf. vitiensis (Seem.) Fader (2102)ConnaraceaeAgelaea borneensis (Hook. f.) Merr. (491)Connarus culionensis Merr. (686)Connarus semidecandrus Jack (623)Ellipanthus tomentosus Kurz (396)CrypteroniaceaeCrypteronia cumingii (Planch.) Planch. ex Endl. (1693)CunoniaceaeWeinmannia cf. hutchinsonii Merr. (130)DatiscaceaeOctomeles sumatrana Miq.DilleniaceaeDillenia megalantha Merr. (2007) VUDillenia philippinensis Rolfe VUTetracera fagifolia Bl. (674)DioscoreaceaeDioscorea hispida Dennst.DipterocarpaceaeAnisoptera thurifera Foxw. ssp. thurifera (353)Dipterocarpus gracilis Bl. (486) CRDipterocarpus validus Bl. CRHopea acuminata Merr. (292) CRHopea malibato Foxw. ex Elm. (20) CRHopea philippinensis Dyer (925) CRHopea plagata (Blco.) Vid. (305) CRParashorea malaanonan (Blco.) Merr. (267) CRShorea almon Foxw. (430) CRShorea assamica Dyer forma philippinensis (Brandis) Sym. (269) CRShorea astylosa Foxw. (1796) CRShorea cf. hopeifolia (Heim) Sym. (2110) CRShorea contorta Vid. (1001) CRLife formMacPMesPMesPMesPMacPMesPCh herbCh herbCh herbCh herbCh herbst PLPLPLMesPMesPMesPMacPMesPMesPst PLPLMacPMacPMacPMacPMacPMacPMacPMacPMacPMacPMacPMakPMacP

1288SpermatophytaShorea falciferoides Foxw. ssp. falciferoides (290) CRShorea guiso (Blco) Bl. (384) CRShorea palosapis (Blco) Merr. (263) CRShorea polysperma (Blco) Merr. (297) CRVatica mangachapui Blco. (528) ENEbenaceaeDiospyros blancoi A. DC. (163) VUDiospyros cf. nitida Merr. (1901)Diospyros curranii Merr. (1631)Diospyros multibracteata Merr. (598)Diospyros pilosanthera Blco.Diospyros pyrrhocarpa Miq. (385)ElaeagnaceaeElaeagnus triflora Roxb. var. triflora (412)ElaeocarpaceaeElaeocarpus cumingii Turcz. (1123)EuphorbiaceaeAcalypha amentacea Roxb. (254)Antidesma digitaliforme Tul. (371)Antidesma nitidum Tul. (268)Antidesma tomentosum Bl. (919)Aporosa benthamiana Hook. f. (573)Baccaurea tetrandra (Baill.) Mu¨ll. Arg. (360)Bridelia glauca Bl. (233)Claoxylon brachyandrum Pax & K. Hoffm. (379)Cleistanthus cf. glaber Airy Shaw (628)Cleistanthus sumatranus (Miq.) Mu¨ll. Arg. (396)Codiaeum luzonicum Merr.Croton cascarilloides Raeusch. (205)Drypetes cf. megacarpa (Bl.) Pax & Hoffm. (374)Drypetes longifolia (Merr.) Pax et Hoffm. (372)Glochidion rubrum Bl. (715)Macaranga caudatifolia Elm. (735) VUMacaranga grandifolia (Blcol.) Merr. VUMacaranga hispida (Bl.) Muell.-Arg.Macaranga tanarius (L.) Muell.-Arg.Mallotus cf. paniculatus (Lam.) Muell.-Arg. (330)Mallotus floribundus (Bl.) Muell.-Arg. (228)Mallotus lackeyi Elm. (1800)Mallotus philippensis (Lam.) Muell.-Arg. (302)Neotrewia cumingii (Muell.-Arg.) Pax & Hoffm. (343)Omalanthus populneus (Geisel.) PaxPhyllanthus leytensis Elm. (250)Suregada glomerulata (Hassk.) Jones (287)FagaceaeLithocarpus buddii (Merr.) A. Camus (15)Lithocarpus caudatifolia (Merr.) Rehd. (555)Lithocarpus coopertus (Blco) Rehd. (387)Life formMacPMacPMacPMacPMacPMacPMiPMesPMiPMesPMesPdPL frutMesPNPNPMiPMicPMiPMesPMesPMesPMesPMesPMiPNPMiPMiPMiPMiPMesPMiPMesPMesPMesPMesPMesPMesPMiPCh frutNPMacPMesPMesP

1289SpermatophytaFlacourtiaceaeCasearia cf. mindanaensis Merr. (1675)Casearia grewiaefolia Vent. var. gelonioides (Bl.) Sleum. (794)Flacourtia cf. indica (Burm. f.) Merr. (378)Osmelia philippina (Turcz.) Benth. (352)FlagellariaceaeFlagellaria indica L. (1128)GesneriaceaeCyrtandra angularis Elm. (2212)Cyrtandra glaucescens Kranzl. (960)Monophyllaea merrilliana Kranzl. (2027)Rhynchoglossum obliquum Bl.GnetaceaeGnetum gnemon L. var. gnemon (375)Gnetum latifolium Bl.HamamelidaceaeSycopsis dunnii Hemsl. (739)HernandiaceaeIlligera megaptera Merr. (721)IcacinaceaeGomphandra cumingiana (Miers) F.-Vill. (1118)Gonocaryum calleryanum (Baill.) Becc. (700)Miquelia celebica Bl.Phytocrine macrophylla (Bl.) Bl. var. macrophyllaPlatea excelsa Bl. var. borneensis (Heine) Sleum. (1217)IxonanthaceaeIxonanthes petiolaris Bl.JuglandaceaeEngelhardtia serrata Bl. (411)LamiaceaeGomphostemma javanicum (Bl.) Bth. (285)LauraceaeActinodaphne apoensis Merr. (1083)Actinodaphne bicolor (Elm.) Merr.Actinodaphne cf. multiflora Benth. (833)Caryodaphnopsis tonkinensis (Lec.) Shaw (441)Cinnamomum mercadoi Vid. (468) VUEndiandra coriacea Merr. (1883)Litsea garciae Vid. (478)Litsea leytensis Merr. (805) VUNeolitsea cf. vidallii Merr. (272) VULife formPMesPMesPMesPel PLCh herbCh herbCh herbCh herbMesPPLMesPPLMesPMesPPLPLMesPMesPMacPCh herbMesPMesPMesPMesPMacPMesPMesPMesPMiP

1290SpermatophytaLeeaceaeLeea aculeata Bl. ex Spreng.Leea guineensis G. Don (255)Leea quadrifida Merr. (546)LeguminosaeAfzelia rhomboidea (Blco.) Vid. VUAlbizia procera (Roxb.) Benth.Albizia saponaria (Lour.) Bl. ex Miq.Archidendron clypearia var. casai (Blco.) I.C. Nielsen (1082)Archidendron pauciflorum (Benth.) Nielsen (1852)Archidendron scutiferum (Blco.) I.C. Nielsen (323)Bauhinia integrifolia Roxb. ssp. cumingiana (Benth.) K. & S.S. Larsen (364)Dalbergia cf. mimosella (Blco) Prain (1435)Desmodium laxum DC. (820)Entada phaseoloides (L.) Merr.Erythrina subumbrans (Hassk.) Merr.Euchresta horsfieldii (Lesch.) Benn. (847)Kingiodendron alternifolium (Elm.) Merr. & Rolfe (357)Ormosia calavensis AzaolaPterocarpus indicus Willd. VUWallaceodendron celebicum Koord. (395)LiliaceaeDianella ensifolia (L.) DC.LoganiaceaeFagraea auriculata Jack ssp. auriculata (851)Fagraea racemosa Jack ex Wall.Strychnos luzoniensis Elm. (748)MagnoliaceaeMagnolia liliifera (L.) Baill. var. angatensis (719)MarantaceaeDonax cannaeformis (Forst. f.) K. Schum. (1006)MarattiaceaeAngiopteris evecta (Forst.) Hoffm. (1188)Marattia pellucida Presl (1444)MelastomataceaeMemecylon paniculatum Jack (311)MeliaceaeAglaia argentea Bl. (642)Aglaia costata Merr. (275) VUAglaia elliptica Bl. (1295)Aglaia luzoniensis (Vid.) Merr. & Rolfe (511)Aphanamixis polystachia (Wall.) R.N. Parker (941)Chisocheton ceramicus (Miq.) C. DC.) (1184)Chisocheton cumingianus (C. DC.) Harms (211)Chisocheton pentandrus (Blco.) Merr. (753)Life formMipMiPMiPMesPMacPMesPMesPMiPMesPPLMesPCh herbel PLMacPCh herbMacPMesPMesPMacPCh herbst PLMiPel PLMesPMiPNP herbCh herbMiPMesPMesPMesPMiPMesPMesPMesPMesP

1291SpermatophytaDysoxylum arborescens (Bl.) Miq. (664)Dysoxylum cumingianum C. DC. (316)Reinwardtiodendron humile (Hassk.) Mabb. (965)Toona calantas Merr. & Rolfe (918)Vavaea amicorum Benth. (273)Walsura cf. pinnata Hassk. (2082)MenispermaceaeArcangelisia flava (L.) Merr. (1798)MonimiaceaeMatthaea pubescens Merr. (139)MoraceaeFicus aurita Bl. (210)Artocarpus blancoi (Elm.) Merr. (1701) VUArtocarpus elastica Reinw. ex Bl. (697)Ficus balete Merr. (v)Ficus benjamina L. (1075)Ficus cumingii Miq. var. angustissima (Merr.) Corner (778)Ficus fistulosa Reinw. ex Bl. (1307)Ficus heteropoda Miq. (425)Ficus odorata (Blco.) Merr.Ficus pedunculosa Miq. (1780)Ficus pseudopalma Blco.Ficus punctata Thunb. (406)Ficus ribes Reinw. ex Bl. (405)Ficus ruficaulis Merr.Ficus subulata Bl. (646, 1966)Ficus ulmifolia Lam. (451) VUMaclura cochinchinensis (Lour.) Corner (1417)Streblus ilicifolia (Vid.) Corner (1700)Streblus macrophyllus Bl. (216, 335, 613)MyristicaceaeEndocomia macrocoma (Miq.) W.J.J. de Wilde subsp. prainii(King) W.J.J.de Wilde (479)Gymnacranthera farquhariana (Hook. f. & Th) Warb. var.paniculata (A. DC.) R. Schouten (541)Horsfieldia cf. costulata (Miq.) Warb. (2002)Knema glomerata (Blco.) Merr. (641)Knema stellata Merr. (1481)Myristica cf. frugifera W. J. J. de Wilde (743) VUMyristica cf. philippensis Lam. VUMyristica simiarum A. DC cf ssp. simiarum (417)MyrsinaceaeArdisia pardalina Mez. (815)Ardisia squamulosa Presl (204) VUMaesa denticulata Mez (241)MyrtaceaeAcmena acuminatissima (Bl.) Merr. & Perry (503)Life formMesPMesPMesPMacPNPMesPel PLMiPMiPMacPMacPMacPMacPMesPMiPMiPMesPMiPNPr PLMiPMesPPLMesPd PLMesPMesPMesPMesPMesPMesPMesPMesPMesPMesPCh frutCh frutMiPMacP

1292SpermatophytaSyzygium cf. densinervium (Merr.) Merr. (749)Syzygium cf. xanthophyllum (C.B. Rob.) Merr.Syzygium cumini (L.) SkeelTristaniopsis decorticata (Merr.) P.G. Wilson& J.T. Waterh. (142) VUTristaniopsis micrantha (Merr.) P.G. Wilson & J.T. Waterh. (301)Xanthostemon verdugonianus Naves (2209) VUOchnaceaeGomphia serrata (Gaertn.) KanisOlacaceaeErythropalum scandens Bl. (780)Strombosia philippinensis (Baill.) Rolfe (380)OleaceaeOlea borneensis Boerl. (306)OpiliaceaeChampereia manillana (Bl.) Merr. (100)Melientha suavis Pierre ssp. suavis (366)OrchidaceaeCalanthe triplicata (Willem.) AmesCeratostylis retisquama Rchb. f.B143Cymbidium aliciae Quis. (880)Eulophia zollingeri (Reichb.f.) J.J.SmithGrammatophyllum multiflorum var. tigrinum Lindley.Lepidogyne longifolia (Bl.) Bl.Liparis wenzelii AmesPhalaenopsis hieroglyphica (Rchb. f.) SweetRobiquetia cf. compressa Schltr.Trichoglottis latisepala AmesTrichoglottis rosea (Lindl.) Ames (1055)PandanaceaeFreycinetia cf. philippinensis Hemsl. (1353)Freycinetia cumingiana Gaudich. (388, 1234)Freycinetia multiflora Merr. (1130)Freycinetia vidalii Hemsl. (1352)Freycinetia membranifolia Elm. (955)PentaphragmataceaePentaphragma grandiflorum Kurz (457, 458, 1407)PiperaceaePiper abbreviatum Opiz (638)Piper halconense C. CD.Piper toppingii C. CD. (654, 1143)Piper viminale Opiz (1205)PittosporaceaePittosporum resiniferum Hemsl. (448)Life formMesPMesPMesPMesPMesPMacPMesPel PLMesPMesPMesPMesPG rhizEEG rhizEG rhizG rhizEEEEr PLr PLr PLr PLr PLCh herbst PLst PLst PLst PLMesp

1293SpermatophytaPoaceaeBambusa spinosa Roxb.Dinochloa cf. pubiramea GambleDinochloa cf. scandens (Bl.) O. Ktze.PodocarpaceaePodocarpus rumphii Bl. (1520)PolygalaceaePolygala venenosa Juss. ex Poir. (284, 2011)Xanthophyllum vitellinum (Bl.) Dietr. (992)ProteaceaeHelicia graciliflora Merr. (1154)Helicia loranthoides Presl. (1079)Helicia robusta (Roxb.) R. Br. ex Wall. (588)RanunculaceaeClematis javana DC. (159, 1997)RhamnaceaeVentilago dichotoma (Blco.) Merr. (723)Ziziphus angustifolius (Miq.) Hatusima ex Steenis (488)Ziziphus crebrivenosa C.B. Rob. (492, 661)RhizophoraceaeGynotroches axillaris Bl. (1538)RosaceaePrunus arborea (Bl.) Kalkm. var. arborea (1624)Prunus cf. fragrans (Elm.) Kalkm. (795)Prunus grisea (Bl.) Kalkm. var. grisea (71, 490)Rubus fraxinifolius Poiret (2017)RubiaceaeBoholia nematostylis Merr. (1919)Canthium gynochthodes Baill. (563)Diodia ocynifolia (Willd.) Brem. (1424)Diplospora cf. fasciculiflora Elm. (663)Dolicholobium philippinense Trenteuse (260)Greeniopsis multiflora (Elm.) Merr. (279)Hedyotis baruensis (Miq.) Val. ex Merr. (329)Hypobathrum purpureum (Elm.) Merr. (1507)Ixora bartlingii Elm. (1060)Ixora cf. cumingiana Vidal (509)Ixora cf. macrophylla Bartl. (207)Ixora longistipula Merr. (1122)Ixora macrophylla Barth.Ixora salicifolia (Bl.) DC. (288)Lasianthus cf. obliquinerva Merr. (701)Morinda bracteata Roxb. (326)Mussaenda philippica A. Rich.Mussaenda vidallii Elm. (129)Mycetia javanica (Bl.) Korth. (258)Nauclea subdita (Korth.) Stend. (1958)Life formMesPPLPLMacPCh herbMesPMiPMesPMiPPLPLMesPd PLMacPMesPMesPMesPd PL suffCh herbMesPPLMiPMiPMesPCh herbMesPMipMiPMiPMiPMiPNPMiPMiPMiPMiPCh suffMiP

1294SpermatophytaLife formNeonauclea formicaria (Elm.) Merr. (793)MiPNeonauclea lanceolata (Bl.) Merr. subsp. gracilis (Vidal) Ridsdale (402)MesPPraravinia cf. mindanensis (Elm.) Brem. (289)NPPsychotria cf. ixoroides Bartl. ex DC. (515)st PLPsychotria membranifolia Bartl. ex DC. (257)NPTarenna cumingiana (Vid.) Elm. (464)MesPTarrenoidea wallichii (Hook. f.) D.D.Tirvengadum & C. Sastre (307)MesPTimonius arboreus Merr. (248)MiPUncaria cf. perrottetii (A. Rich.) Merr. (325)el PLUncaria lanosa Wall. f. philippinensis (Elm.) Ridsd. (900)el PLUncaria longiflora (Poir.) Merr. (1300)el PLWendlandia luzoniensis DC. (444)MesPXanthophytum fruticulosum Reinw. ex Bl. (1005)NPRutaceaeClausena anisum olens (Blco.) Merr. (605) NPLunasia amara Blco. (158)NPMicromelum compressum (Blco.) Merr. (771)NPSeverinia disticha (Blco) Merr. (398)NPSapindaceaeAllophyllus cobbe (L.) Raeuschel (823)MesPCubilia cubili (Blco.) Adelh. (586)MacPDictyoneura acuminata Bl. ssp. acuminata (246)MesPDimocarpus fumatus (Bl.) Leenhouts ssp. philippinensis Leenhouts (72)MesPEuphorianthus obtusatus Radlk. ex Koord. (1641)MesPGanophyllum falcatum Bl. (1212)MesPGuioa cf. diplopetala (Hassk.) Radlk. (1104)MesPHarpullia cupanioides Roxb. (212)MesPLepisanthes fruticosa (Roxb.) Leenh. (933)MesPNephelium cf. ramboutan ake (Labill.) Leenh. (442) MesPParanephelium cf. xestophyllum Miq. (727)MesPPometia pinnata Forst. (578, 1546)MesPSapotaceaePalaquium philippense (Perr.) C. B. Rob. (443) VUMacPPlanchonella mindanaensis Clemens (1126)MacPPouteria firma (Miq.) Baehni (1237)MacPSaxifragaceaeDichroa philippinensis Schltr.Ch frutPolyosma integrifolia Bl. (1219)NPDichroa fibrifuga (807)SimaroubaceaePicrasma javanica Bl. (218)MesPSolanaceaeSolanum anisophyllum Elm. (225)Ch herbSolanum ferox L. (282)Ch herbSonneratiaceaeDuabanga moluccana Bl.MacP

1295SpermatophytaStaphyleaceaeTurpinia borneensis (Merr. & Perry) B.L. Linden (1802)SterculiaceaeHeritiera sylvatica Vidal (1768)Pterocymbium tinctorium (Blco.) Merr. (345)Pterospermum diversifolium Bl. (270)Pterospermum elongatum Korth. (434)Pterospermum obliquum Blco. (120)Sterculia multistipularis Elm. (251)Sterculia oblongata R. Br. (678)Sterculia philippinensis Merr. (1898)Sterculia stipulata Korth. var. jagorii (Warb.) TantraSymplocaceaeSymplocos cochinchinensis(Lour.) Moore var. cochinchinensis (1954)TaccaceaeTacca palmata Bl. (303)TheaceaeEurya acuminata DC. (1314)Ternstroemia philippinensis Merr. var. philippinensis (1491)ThymelaeaceaeAquilaria cumingiana (Decn) Ridl. (300)Phaleria perrottetiana (Dcne) F.-Vill. (160)TiliaceaeColona serratifolia Cav. (626)Diplodiscus paniculatus Turcz. (271) VUUlmaceaeCeltis cf. philippinensis BlancoGironniera celtidifolia Gaudich. (238)Trema orientalis (L.) Bl. (2202)UrticaceaeCypholophus moluccanus (Bl.) Miq.Leucosyke capitellata (Poir.) Wedd. (242)Maoutia setosa Wedd.Villebrunea rubescens (Bl.) Bl. (324)Villebrunea trinervis Wedd. (733)VerbenaceaeClerodendrum villosum Bl.Teijsmanniodendron pteropodum (Miq.) Bakh. (157)Vitex parviflora Juss. (1837)Vitex turczaninowii Merr. (705)Premna odorata Blco. (633)Life formMesPMesPMacPMesPMesPMesPMiPMesPMesPMesPMicPG rhizNPMesPNPCh suffMesPMesPMesPMiPMesPCh frutMiPNPMesPMesPNPMesPMacPMacPMesP

1296II.PtaridophytaAspidiaceaeDidymochlaena cf. truncatula (Sw.) J. Sm. (2056)AspleniaceaeAsplenium nidus L. (1902)Asplenium tenerum Forst. (2096)AthyriaceaeDiplazium asperum (Bl.) Milde (1809)Diplazium esculentum (Retz.) Sw. (1846)CyatheaceaeCyathea cf. contaminans (Hook.) Copel.DavalliaceaeDavallia solida (G. Forst.) Sw. (1462)Davallia trichomanoides Bl. var. lorrainii (Hance) Holttum (222)HymenophyllaceaeTrichomanes javanicum Bl. (1042)LindsaeaceaeLindsaea lucida Bl. ssp. lucida (533)Sphenomeris chinensis (L.) MaxonTapeinidium pinnatum (Cav.) C.Chr. (1267)LomariopsidaceaeBolbitis cf. guoyana (Gaudich.) Ching (2016)Bolbitis guoyana (Gaudich.) ChingBolbitis heteroclita (Presl) Ching (1049)Lomogramma cf. copelandii Holttum (1851)Lomogramma copelandii HolttumTeratophyllum arthropteroides (Christ) Holttum (2084)Teratophyllum cf. articulatum (J. Sm. ex Fèe) Mett. (516)OsmundaceaeOsmunda banksiaefolia (Pr.) Kuhn (1261, 1392)PolypodiaceaeDrynaria quercifolia (L.) J. SmLeptochilus cf. decurrens Bl.Microsorum cf. longissimum J. Sm. ex Fe´e (964)Microsorum membranifolium (R. Br.) ChingMicrosorum punctatum (L.) Copel. (1821)Microsorum scolopendria (Burm. f.) Copel. (1445)Pyrrosia cf. lanceolata (L.) FarwellMicrosorum plukenetii (Presl) M.G. Price (1860)PteridaceaePteris cf. pellucida PreslPteris ensiformis Burm. f. (1806)Pteris longipinnula Wall. (334)SchizaeaceaeLygodium auriculatum (Willd.) Alst. et Holtt. (1974)Lygodium circinnatum (Burm. f.) Sw. (1603)Life formCh herbCh herbCh herbCh herbCh herbMesPCh herbCh herbCh herbCh herbCh herbCh herbCh herbCh herbr PLr PLr PLCh herbCh herbCh herbECh herbCh herbCh herbCh herbCh herbPLCh herbCh herbCh herbst PLst PL

1297PtaridophytaSelaginellaceaeSelaginella cf. involvens (Sw.) Spring (856)Selaginella cf. springiana Alderw. (1526)Selaginella cupressina (Willd.) Spring (745)Selaginella engleri Hieron. (1011)TaenitidaceaeTaenitis blechnoides (Willd.) Sw. (1091)Tectaria groupCtenitis cf. silvatica Holttum (939)Cyclopeltis crenata (Fée) C. Chr. (1807)Pleocnemia cf. presliana Holttum (1849)Pleocnemia irregularis (Presl) Holttum (1007)Tectaria crenata Cav. (1301)ThelypteridaceaeCyclosorus sumatranus (v. Ald. v. Ros.) ChingPneumatopteris laevis (Mett.) Holttum (1812)Pronephrium · xiphioides (Christ) Holttum (498)Pseudophegopteris paludosa (Bl.) Ching (2093)Pronephrium granulosum (Presl) Holtt. (997)Life formCh frutCh frutCh frutCh frutCh herbCh herbCh herbCh herbCh herbCh herbCh herbCh herbCh herbCh herbCh herbReferencesAnonymous 1993. Flora of Taiwan Vol. 3, Angiosperms, Dicotyledons, , 2nd edn. EditorialCommittee of the Flora of Taiwan, Department of Botany, National Taiwan University, Taipei,Taiwan, ROC.Anonymous 1994. Flora of Taiwan Vol. 1, Pteridophyta, Gymnospermae, 2nd edn. EditorialCommittee of the Flora of Taiwan, Department of Botany, National Taiwan University, Taipei,Taiwan, ROC.Anonymous 1996. Flora of Taiwan Vol. 2, Angiospermae, 2nd edn. Editorial Committee of theFlora of Taiwan, Department of Botany, National Taiwan University, Taipei, Taiwan, ROC.Aragones E.G. Jr. 1991. Vegetation-soil pattern along altitudinal gradient in the western slopes ofMt. Banahaw, Luzon, Philippines: I. The forest communities and changes in forest compositionwith altitude. Sylvatrop 1(1): 15–45.Ashton P.S. 1993. Philippine phytogeography. Asia Life Sciences 2(1): 1–8.Asio V. 1996. Characteristics, weathering, formation and degradation of soils from volcanic rocksin Leyte, Philippines. Hohenheimer Bodenkundliche Hefte 33 Universität Hohenheim Institutfu¨r Bodenkunde und Standortslehre, Stuttgart, 209 pp.Barrera A., Aristorenas I. and Tingzon J.A. 1954. Soil Survey of Leyte Province, Philippines. SoilSurvey Report No. 18, Bureau of Printing, Manila, 103 pp.Bremer H. 1995. Boden und Relief in den Tropen: Grundvorstellungen und Datenbank. Gebru¨derBornträger, Berlin, Stuttgart, 324 pp.Bremer H. 1999. Die Tropen – Geographische Synthese einer fremden Welt im Umbruch. Gebru¨derBornträger, Berlin, Stuttgart, 428 pp.Brooks T.M., Mittermeier R.A., Mittermeier C.G., da Fonseca G.A.B., Rylands A.B., KonstantW.R., Flick P., Pilgrim J., Oldfield S., Magin G. and Hilton-Taylor C. 2002. Habitat loss andextinction in the hotspots of biodiversity. Conservation Biology 16(4): 909–923.

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