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Woody plant communities in the Philippine teak forest landscape along Verde
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DOI: 10.13057/biodiv/d201111

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BIODIVERSITAS ISSN: 1412-033X
Volume 20, Number 11, November 2019 E-ISSN: 2085-4722
Pages: 3189-3198 DOI: 10.13057/biodiv/d201111

Woody plant communities in the Philippine teak forest landscape along

Verde Island Passage, Batangas, Luzon, Philippines

ANACLETO M. CARINGAL1,2, INOCENCIO E. BUOT JR.1,3,4,♥, ELAINE LOREEN C. VILLANUEVA3

1
School of Environmental Science and Management, University of the Philippines Los Baňos. College Laguna 4031, Philippines
email: iebuot@up.edu.ph
2
Program for Tropical Forestry, College of Agriculture and Forestry, Batangas State University-Lobo. Lobo Batangas 4229, Philippines
3
Institute of Biological Sciences, College of Arts and Sciences, University of the Philippines Los Baňos. College Laguna 4031, Philippines
4
Faculty of Management and Development Studies, University of the Philippines Open University. Los Baňos Laguna, Philippines

Manuscript received: 17 December 2018. Revision accepted: 14 October 2019.

Abstract. Caringal AM, Buot IE Jr, Villanueva ELC. 2019. Woody plant communities in the Philippine teak forest landscape along
Verde Island Passage, Batangas, Luzon, Philippines. Biodiversitas 20: 3189-3198. The study described the communities of woody
plants in a semi-evergreen tropical forest dominated by endemic Philippine teak (Tectona philippinensis Benth. & Hook. f, Lamiaceae)
and determined the local environmental factors affecting the distribution of dominant woody species across the Philippine teak forest
(PTF) landscape. The Philippine teak is a Critically Endangered species according to the International Union for the Conservation of
Nature (IUCN). Quantitative dominance analysis of arboreal vegetation data (basal area and relative dominance) from 24 (20m x 20m)
sampling plots generated the cluster dendrogram, while Canonical Correspondence Analysis (CCA) was performed for 47 woody
species and habitat-environment variables. As a result, four vegetation zones were named: (I) pure stand of Tectona philippinensis, (II)
mixed T. philippinensis-Garuga floribunda-Terminalia polyantha, (III) mixed Celtis latifolia-T. philippinensis and (IV) mixed
Tamarindus indica-Xylocarpus rumphii. These zones were governed more by physiographic (altitude ) and edaphic influences of
predominantly agro-coastal landscapes. The environmental variables were essential in verifying not only the association of dominant
tree flora in the local landscape but also provide insight for forest management zoning and the ecological requirements of the tree
species for in-situ and ex-situ biodiversity conservation.

Keywords: Endemic species, Philippine teak, Tectona philippinensis, Verde Island Passage

INTRODUCTION Tan and Rojo 1988; Madulid 1994; Fernando et al. 2008).
The Philippine teak forest (PTF), however, has not yet been
The tropical forests of Southeast Asia is distinct in widely studied among the unique ecotypes in the
terms of floristic composition and diversity due to complex Philippines.
biogeography and environmental heterogeneity (Whitmore In Batangas Province along the Verde Island Passage,
1975, 1988; Newbery et al. 1992; Buot and Okitsu 1998; the forest with Philippine teak (Tectona philippinensis
Newbery et al. 1999; Belonias and Aguilar 2004; Buot and Benth. & Hook. f, Lamiaceae) occurs as fragments across
Osumi 2011; Heng et al. 2017). Forest vegetation has a the agro-coastal landscape at low altitudes (05-200 m asl.
subtle response to environmental influences, resulting in particularly in the municipalities of Lobo, San Juan, and
variation in species composition and structure, and serve as Taysan and Batangas City where the natural tree
bio-indicators of landscape productivity and human populations show spectacular flowering episodes preceding
disturbances (Buot 2007; 2008; Sopsop and Buot 2013; a long summer (Agoo and Oyong 2008; Caringal et al.
Nguyen et al. 2015; Van and Cochard 2017). Thus, it was 2015). Tectona philippinensis was first described by British
always of interest and value to investigate the plant botanists George Bentham and Joseph D. Hooker (1876)
communities in this landscape (Whittaker 1960, 1965; based on the specimens deposited at London’s Kew
Mueller-Dombois and Ellenberg 1974; Aragones 1991; Herbarium which was collected by English naturalist Hugh
Villanueva and Buot 2018). Cuming (1432) from the shores of Batangas, Luzon.
In the Philippines, there is the existence of diverse Because of its limited and fragmented distribution, the
natural forests known as ecotypes or formation species is of special interest to phytogeographers in the
characterized by distinct species composition and structure, Malesian Region (Madulid and Agoo 1990). Based on the
which were classified based on the dominant tree elements assessment criteria of the International Union for the
(Department of Environment and Natural Resources- Conservation of Nature (IUCN), T. philippinensis has been
United Nations Environment Programme, 1997; Baguinon classified as Critically Endangered (Madulid et al. 2008).
2000; Malayang 2001; Pulhin 2002). These formations Given the narrow distribution and current conservation
ranged from sea level up to 2,954 m altitudes and were the status of Tectona philippinensis, it is therefore important to
focus of ecological classification since the 1900s (Whitford study the ecological aspect of the PTF to provide in-depth
1909, 1911; Whitmore 1984; Cadiz 1986; Fernando 1988; and significant information about this unique ecotype in the
3190 B I O D I V E R S I T A S 20 (11): 3189-3198, November 2019

Philippines. As there were fewer local landscape-level (Pleiocene-Pleistocene) and San Juan Quartz Diorite
studies about the Philippine teak, this study was undertaken (Lower Miocene) where the underlying geography is
to reveal the dominant floristic elements according to plant shallow, soft and porous reef limestone (Pleistocene)
community patterns of the Philippine teak forest. Using the including bushes of corals, shells, algae structure and
data gathered from the vegetation surveys in the Philippine recently alluvium formation (Bureau of Soils 1987; Bureau
teak forest, the researchers used classification analysis of Mines and Geo-Sciences-Mindoro Resources Limited
(Cluster Analysis) in identifying the plant communities in Gold Philippines Inc 1981; Caringal 2007; Comprehensive
the study area. Moreover, ordination analysis (Canonical Land Use Plan of Lobo, Batangas 2012; Cox 2010;
Corresponding Analysis) was also utilized in investigating Mindoro Resources Limited 2004). The mean annual
the influence of the environment and anthropogenic factors precipitation (1980-2016) is 151.89 mm in Batangas
on the species distribution in the Philippine teak forests. The Province (Figure 2). The precipitation in the province
results can be used in forest management zoning particularly reaches is at its highest records in July (331mm). Low
in locally and nationally important ridge-to-reef biodiversity rainfall (27-32mm) was experienced during summer,
conservation corridors of Verde Island Passage. January to April. Annual mean temperature (AMT) was
27.36°C ranging from 26°C-29°C (Figure 2).

MATERIALS AND METHODS Procedures

Forest vegetation survey
Study area Fieldworks were conducted from October 2016-
The vegetation survey was conducted along Batangas February 2017 and resulted in a total of 24 (20m x 20m)
Province, particularly in the areas of Batangas City, vegetation plots (=0.96ha). The sampling locations were
municipality of Lobo, and Isla Verde (08-321 m asl, identified before successive field surveys of the Tectona
13°30’-13°40’ North latitudes and 121°05’-121°15’East philippinensis forests. The plots were purposively selected
longitudes) in southwestern Luzon, Philippines (Figure 1). based on (i) the quality of trees and associated plants and
Twenty-four (24) 20m x 20m vegetation plots were taken (ii) the presence of human-related disturbances or
from Isla Verde to the mainland (56.48 km connectivity), expansion activities that may affect the forest structure. Six
which have gentle to undulating, rolling to moderately vegetation plots were located in the coastal zone (including
steep slopes (0.10 to 51.99%). The plots were located for 1 island coast), three plots in a narrow inland valley, and 15
some distances (58.5m to 6225.28m) along the coastal cliff plots in inland ridges-all lying apart west to east from north
towards inland ridges and narrow valley facing the sea of to south directions (Figure 1). Vegetation assessment was
Verde Island Passage (Figure 1). The landscape consists of done following the techniques of Mueller-Dombois and
volcanic agglomerates from various ages such as Talahib Ellenberg (1974).
Andesite (Upper Oligocene-Miocene), Lobo agglomerate

Figure 1. The Philippine teak forest landscape along the Verde Island Passage, Batangas Luzon, Philippines. The study plots were
shown as small 1-24 black squares from an island to coastal-mainland of Batangas Province, Philippines
 CARINGAL et al. – Trees of Philippine teak forest 3191

Dominance analysis
The dominant species were identified to further
describe each of the plant community that was identified
from the cluster analysis. In this study, the relative basal
area (RBA) of the woody species was used as a measure of
dominance. These were quantitatively identified using the
of Ohsawa (1984):

Where:
d : The deviation
xi : The actual percent share [in this case, the RBA] of
the top species (T), i.e, the top dominant in the one-
dominant model, or the two top dominants in the two-
Figure 2. The 36-Year (1980-2016) climogram showing the dominant model and so on
monthly average precipitation and temperature for Batangas x : The ideal percent share based on the
Province, Philippines (Data retrieved from Synoptic Station aforementioned model
Records: Ambulong, Batangas. Philippine Atmospheric, xj : The percent share of the remaining species (U)
Geophysical and Astronomical Services Administration
(PAGASA), Diliman, Quezon City, Philippines)
N : The total number of species.

Consequently, each plant community (cluster) was

named after the dominant and co-dominant tree flora, i.e.
In each plot, the diameter at breast height (DBH) of species with the highest RBA values. The number of the
trees (at least ≥10cm) was measured with a diameter tape dominant species that represent the plant communities was
based on the formula above.
at 1.3 m from the ground while heights were estimated
visually following Erenso et al. (2014). Crown dimensions
were measured by a meter tape following Almazol and Proximity analysis for physiographic and anthropogenic
Quintana (2009). Altitudes and coordinates of all study factors
plots were taken using a Garmin 76Csx Global Positioning Proximity analysis of the Philippine teak plots to the
System (GPS) receiver. various physiographic regime such as altitude (m asl.) and
Woody plants were identified with the help of local distances (m) to or from the streams, coast or shoreline and
botanists and verified with photographs of plants from the anthropogenic disturbances (e.g. roads, built-up areas,
field and herbarium vouchers lodged at the Batangas State settlements, farms) was undertaken by an ArcGIS
University Herbarium (BatStateU-H) and Plant Biology specialist. This computed the approximate straight line
Division Herbarium of the Institute of Biological Sciences, distance (measured in meters) between a plot and any
University of the Philippines at Los Baňos (UPLB). spatial object, given a UTM 51N projected coordinate
Species nomenclature followed the Revised Lexicon of system and a WGS 1984 geographic projection. The stream
Philippine Trees (Rojo 1999), An Enumeration of network used for the analysis was topographically derived
Philippine Flowering Plants (Merrill 1923-1926), from NAMRIA's 5m spatial resolution IfSAR DEM,
Flowering Plants and Ferns of Mt. Makiling (Fernando et provided that first-order streams have a minimum
al. 2004) and Dictionary of Philippine Plant Names catchment area of 20,000m². The road network is from
(Madulid 2001). Open Street Maps, and built-up areas were isolated from
the latest available land cover map of NAMRIA. The
results of the GIS-based approximate analysis, however,
Data analysis
was in some aspects validated with the actual observation
Cluster analysis
in the studied plots, hence there were some data fine-
To identify the plant communities in the forest
tunings.
landscape, the basal area (BA) values of the woody species
were subjected to cluster analysis using the unweighted
Edaphic environment analysis
pair group method with arithmetic mean (UPGMA)
These are the results of laboratory analysis done on
algorithm and Euclidean distance as the index of similarity
basic soil chemical properties such as acidity (pH),
among the plots. The PAleontological STatistical (PAST)
moisture content (%MO), organic matter (% OM) and
software (Hammer et al. 2010) was used to run the analysis
organic carbon (% OC), nitrogen (% N), phosphorus (P
and generate the cluster dendrogram of the plots. The
ppm), and potassium (K ppm). Soil samples in each plot
diagram was used to visualize the representation of the
were collected uniformly following the procedures
clusters of the vegetation plots as plant communities in the
formulated by PCARR (1980). Soil sampling point (25 cm
landscape.
x 25 cm x 30 cm pit) was dug using an auger and spade
3192 B I O D I V E R S I T A S 20 (11): 3189-3198, November 2019

from four corners of each plot and 1 from the middle. At Cluster I. Pure Tectona forest (33-213m asl.): It was
least 1kg soil slices from these 5 points were mixed formed from a broadleaved monsoon forest, turning
thoroughly and cleared with stones, grasses, weeds or other completely deciduous in summer months (January-April)
plant debris. Each 1kg soil sample was sealed in a plastic located in inland ridges (Figure 7), narrow valley and coast.
bag, labeled and were analyzed at the Regional Soils This plant community has 25 woody species with many
Testing Laboratory of the Philippine’s Department of indigenous trees such as Memecylon edule, Terminalia
Agriculture (DA) in Lipa City, Batangas. These polyantha, Vitex parviflora, Sterculia foetida and Hibiscus
quantitative values were identified and used in analyzing tiliaceous. There are many anthropogenic disturbances in
the influence of the environment variables (including soil almost all the plots (Table 2). The tree height were ranged
data) on the formation of the plant communities in the from 4 to-22 m while DBH ranged from 10 to 77.66 cm.
landscape. The maximum basal area for the most dominant species,
i.e. T. philippinensis was 748.01 m2 and with RBA of
Canonical correspondence analysis 72.65%.
To explain the relationship of dominant tree flora in the Cluster II. Mixed Tectona-Garuga-Terminalia forest
Philippine teak forest with the habitat environment factors, (37-321 m asl.): This cluster is a broadleaved semi-
Canonical Correspondence Analysis (CCA) of Ter Braak deciduous forest from inland ridges to the coast composed
(1986) was performed using the statistical software R of other co-dominant species such as Vitex parviflora,
version 3.4.2 (R Core Team, 2013) with the vegan package Gliricidia sepium, Dehassia triandra, Memecylon edule,
(Oksanen et al. 2007). Environmental factors including soil Wrightia pubescens and Scolopia luzonensis. The plots
data, altitude, slope and proximity to anthropogenic were also affected by various anthropogenic disturbances
disturbances were tested (Table 1). These environmental (Table 2). This plant community had 36 woody species
gradients are the basis for visualizing (modelling) and with DBH range from 4.14-60.48 cm and height of 6-23 m.
describing the differential habitat preferences (niches) of The combined total basal area for the three dominant
taxa through an ordination diagram. species was 1,103.98 m2 with T. philippinensis having the
highest RBA of 36.43%.
Cluster III. Mixed Celtis-Tectona (53 m asl.): This is a
RESULTS AND DISCUSSION semi-evergreen forest in a coastal cliff (Plot 5) with very
few (5) and scanty woody species adjoining uphill swidden
Plant communities of the woody plants in the Philippine patches overlooking the coastal road and dense settlement
teak forest landscape below. In this zone, Celtis latifolia is the evergreen
The dendrogram generated from cluster analysis of broadleaved tree associated with deciduous T.
woody species basal area (BA) revealed four plant philippinensis. Tree DBH ranged from 3.81-57.93 cm
communities (Table 2, Figure 3) named according to their while heights were 5-15 m. The combined basal area of the
dominant species: (I) pure Tectona philippinensis (33-213 dominants (C. latifolia and T. philippinensis) was 44.54 m2.
m asl.), (II) mixed Tectona-Garuga-Terminalia (37-321 m The RBA of C. latifolia was 51.51%, while T.
asl.), (III) mixed Celtis-Tectona (53 m asl.) and (IV) mixed philippinensis was 35.54%.
Tamarindus-Xylocarpus (8-209 m asl.) (Figure 4). Cluster IV. Mixed Tamarindus-Xylocarpus (8-209 m
asl.): This is an evergreen forest in ridge valley and beach
coast composed of 12 woody species having DBH that
ranged from 3.81-91.99 cm and heights ranged from 3-
Table 1. Environmental variables used in Canonical 20m. In this plant community, T. philippinensis was
Correspondence Analysis (CCA) overshadowed by most dominant species Tamarindus
indica with a basal area of 161.42 m2 and RBA of 52.79%
Environmental variable Abbreviation and, Xylocarpus rumphii with a basal area of 66.74m2 and
Physiographic Percent slope SLP RBA of 21.83%. In this zone, Terminalia catappa,
Proximity to the sea SEA Thespesia populnea, and Guettarda speciosa are co-
Proximity to the river RIV dominant trees.
Altitude ALT
Environmental factors influencing the distribution of
Edaphic Soil moisture MO woody plants
Organic carbon OC
Organic matter OM
Using Canonical Correspondence Analysis (CCA), the
pH pH distribution of 24 sampling plots (quadrats) and 47 woody
Phosphorus P species over environmental gradients were presented in
Potassium K Figure 5 and Figure 6. The eigenvalue of Axis 1 was 0.531
while Axis 2 was 0.408 with 24.67% total inertia (weighted
Anthropogenic Proximity to built-up areas/ SETL variance) and 42.12% of the variance in the weighted
settlement averages and class totals of the species with respect to
Proximity to swidden or farm FR environmental variables. The CCA biplot showed that
Proximity to road RD habitat environment variables affected the distribution and
dominance of species in the forest.
BIODIVERSITAS CARINGAL et al. – Trees of Philippine teak forest ISSN: 1412-033X
3193
Volume 20, Number 11, November 2019 E-ISSN: 2085-4722
Pages: 3189-3198 DOI: 10.13057/biodiv/d201111
Table 2. Tree communities, distribution of the sample plots and observed human disturbances in the Philippine teak forest landscape

Tree or Observed
woody Altitude Slope Dominant ∑BA RBA Anthropogenic
Tree communities Location
species (m asl.) (%) trees (m2) (%) Disturbances
richness (ADs)*
Cluster I 25 33-213 0.61-86.17 Tectona 748.01 72.65 SCP, PGL, AT, Inland ridges,
(plots: 3, 4, 7, 16, 17,13, philippinensis FLG, Ro, HS valley, coast
20,1, 24, 22, 12)
Cluster II 36 37-321 2.68- T. philippinensis 521.43 36.43 SCP, PGL, AT, Inland ridges,
(plots: 6, 14, 15, 8, 18, 251.99 Terminalia polyantha 395.84 27.66 FLG, Ro, HS coast
9, 10, 21, 11, 19) Garuga floribunda 186.71 13.05

Cluster III 5 53 32.02 Celtis latifolia 26.36 51.51 SCP, PGL, AT, Coastal cliff
(plot: 5) T. philippinensis 18.18 35.54 FLG, Ro, HS

Cluster IV 12 8-209 56.87- Tamarindus indica 161.42 52.79 AT, FLG, Ro, HS Ridge valley
(plots: 2, 23) 97.88 Xylocarpus rumphii 66.74 21.83 and ridge coast
*Notes: Observed Anthropogenic Disturbances (ADs): SCP: Near swidden and crop plantations, e.g. ates, ipil-ipil, mahogany, other
cropping (within 5-20 m distance); PGL: Near pasture or grazing land or the study plots with ipil-ipil source of forage (within 5-20m
distance); AT: Near access trail within 5-20m distance; FLG: The plots with remnant cuts of fuelwood or lumber gathering; Ro: Near
road within 5-100m distance; HS: Near human settlement within 10-100m distance

Figure 3. Dendrogram generated from cluster analysis based on woody species basal area (BA) in the Philippine teak forest landscape
using the UPGMA algorithm with cophenetic correlation of 0.88. At a Euclidean distance between 50-60, four clusters were identified
with corresponding dominant tree species having the highest relative basal area (RBA).
3194 B I O D I V E R S I T A S 20 (11): 3189-3198, November 2019

Figure 4. Photographs of four clusters (zones) of woody plants in the Philippine teak forest landscape representing ridge-to-coast habitats

The result of ordination is complementing the cluster become typically deciduous during summer and extended
analysis which modelled four plant communities (Figure drought. Garuga floribunda was a typical tree of the semi-
3). On the upper portion of the biplot was Cluster II, deciduous forest from 0-900 m altitude in Sierra Madre and
associated with edaphic factors such as increasing level of Palawan, but in Batangas it was a rare co-dominant with
potassium (K), soil moisture (MO) and pH. Cluster III (Plot endemic T. philippinensis. The PTF occurs with beach flora
5 MC) was more sensitive to moisture and pH levels, while such as Celtis latifolia (Cluster III) but overshadowed in
Cluster IV (Plots 2 RV and 23 MC) except with terms of dominance by Xylocarpus rumphii and
phosphorous level was diverged from most of the edaphic Tamarindus indica (Cluster IV). Though T. indica was
vectors and was also influenced by slope gradient (Figure encountered on the island and mainland beach coast or
5, Figure 6). The lower part of the second axis showed that coastal cliff, it has a very low affinity with the beach
Zone I (valley and ridge plots) was greatly influenced by forest’s X. rumphii as the former was more frequent in
physiographic factors such as altitude (ALT) and distance inland valley and ridges.
from the coast (SEA). The vegetation succession or the dynamic of habitat-
However, considering the holocoenotic nature of the environment interaction can be initiated by the interplay of
environment, these seemingly isolating influences of the following factors: [i] floristic composition of a given
certain edaphic or physiographic factors might also be landscape, [ii] ability of a given species to reach the
coupled with other influences such as low to moderate habitat, [iii] eco-physiology and morphology of plant life,
effects of slope (SLP), other edaphic attributes such as [iv] the habitat, and [v] the time that has elapsed following
level of organic matter (OM), organic carbon (OC) and a major disturbance (Mueller-Dombois and Ellenberg
phosphorous (P); anthropogenic disturbances such as roads 2002). The Philippine teak plant communities can be best
(RD), farming (FR) and settlement (SETL) or by seen as a consequence of habitat-environment gradient
combination of all these factors. Interestingly, the sites filtering, as well as habitat specialization, as proposed by
were ordained generally far from riparian influence (RIV), Oliveira et al. (2014) for the “restinga” forest in the
suggesting that PTF is a formation characteristic of the dry Brazilian coast, where more abundant species in “restinga”
woodland environment along Verde Island Passage. forest has niche overlap with other species in at least one
habitat. Closely related species tended to occur in different
Discussion habitats while neighboring trees tended to belong to more
The communities of woody plants have their own distantly related species (Oliveira et al. 2014). This made
distinct characteristics species. There was the pure natural the structure of the ecological species groups to be associated
stand of T. philippinensis in Cluster I while mixed-stand with a combination of habitat factors such as elevation, soil
of T. philippinensis, Terminalia polyantha and Garuga pH, vegetation type and management disturbance regime
floribunda in Cluster II. The forest canopy in these zones and, thus, good indicators (Quimio et al. 2013).
BIODIVERSITAS CARINGAL et al. – Trees of Philippine teak forest ISSN: 1412-033X
3195
Volume 20, Number 11, November 2019 E-ISSN: 2085-4722
Pages: 3189-3198 DOI: 10.13057/biodiv/d201111

Figure 5. The results of ordination analysis in the studied area over 13 environmental variables using Canonical Correspondence
Analysis (CCA) based on relative basal area (RBA) of 47 woody species. The 13 environmental variables were: physiographic-percent
slope (SLP), proximity to the sea (SEA) and river (RIV), and altitude (ALT) above the sea; edaphic-levels of soil moisture (MO),
organic carbon (OC), organic matter (OM), pH, phosphorous (P) and potassium (K) and anthropogenic-proximity to built-up areas,
settlement (L or SETL), swidden or farm (FR) and road (RD). The objects (plots 1-24, green squares) are representing four vegetation
zones across satoyama-satoumi landscape such as mainland ridges (MRs), ridge valleys (RVs), mainland coast (MC) and island coast
(IsC). The four clusters of plots are encircled in red

Figure 6. CCA ordination diagram showing the distribution of the woody species in the biplot (orange dots S1 to Sn…). The figure also
indicated the 24 sampling sites (green squares, unlabeled) and 13 environmental variables influencing the distribution patterns of the
woody species. The Sn are descriptors of 47 woody species under the following genera: Spondias (S1), Wrightia (S2), Garuga (S3),
Bauhinia (S4), Tamarindus (S5), Celtis (S6), Capparis (S7), Crateva (S8), Salacia (S9), Terminalia catappa (S10), T. polyantha (S11),
Cycas (S12), Diospyros (S13), Macaranga (S14), Hibiscus (S15), Melanolepis (S16), Gliricidia (S17), Scolopia (S18), Cratoxylum
(S19), Tectona philippinensis (S20), Vitex (S21), Dehassia (S22), Berrya (S23), Bombax (S24), Colona (S25), Grewia (S26),
Pterocymbium (S27), Pterospermum (S28), Sterculia (S29), Thespesia (S30), Memecylon (S31), Xylocarpus (S32), Acacia (S33),
Albizia (S34), Leucaena (S35), Ficus sumatrana (S36), F. ulmifolia (S37), F. variegata (S38), Antidesma (S39), Canthium (S40),
Guettarda (S41), Mussaenda (S42), Tarrena (S43), Murraya (S44), Dimocarpus (S45), Dodonaea (S46) and Harpaullia (S47).
3196 B I O D I V E R S I T A S 20 (11): 3189-3198, November 2019

Figure 7. The Philippines teak forest (PTF) in inland ridge. The lilac canopies are all individual Tectona philippinensis during mass
flowering on the onset of southwest monsoon in Batangas Province, Philippines

Edaphic and physiographic factors displayed influence make the PTF landscape a very dry one (Whitford 1911).
on the formation of plant communities and species The arid environment of PTF can be also attributed to
distribution patterns. Available edaphic data analysis anomalous macroclimate turning the forest completely
further revealed that PTF occurred in slightly acidic to deciduous, creating canopy gaps and exposing the
acidic soils (pH 5.73); low in moisture (11.88%) but still landscape to direct sunlight during summer and periodic
has an adequate organic matter (6.08%); medium organic prolonged droughts (Madulid and Agoo 1990; Caringal and
carbon content (3.54%), medium amount of phosphorus Makahiya 2000; Caringal et al. 2015; Briones et al. 2018).
(8.37ppm) and extremely high potassium content Under these stressful edaphic-climatic conditions, it was
(360.5ppm). Given such data-based analysis, the PTF can found out that T. philippinensis had already developed
be seen improving soil quality or evolved by sets of the special physio-anatomical features related to persistent
valuable edaphic environment (forest-soil interaction), adaptation in xeric habitat (Hernandez et al. 2016). Edaphic
hence the dominant T. philippinensis can be considered factors such as moisture availability and nutrients directly
among the edaphic-endemic species of the Philippines. The influence species niche differentiation and spatial
Philippine teak forests occurred along coastal hills, littoral distribution of tropical trees at the local-regional scales
cliffs and inland limestone ridges with shallow sedimentary (Engelbrecht et al. 2007). The assembly of tropical tree
rock formation known as Mt. Santiago Limestone communities at plot scales is largely influenced by soils
developed during the Pliocene and Pleistocene period and habitat factors in the landscapes (John et al. 2006).
(Madulid and Agoo 1990; Bureau of Soils 1987; Caringal As a semi-deciduous secondary forest, PTF depends on
et al. 2015). Generally, limestone substrate have shallow the dynamic aspect of physiography as it grows on low
clayey topsoil consisting mainly of porous calcium relief, slightly dissected limestone hills and abyss with
carbonate and sands (Cox 2010), low in water and nutrient slopes ranging from 18 to 50% characterized by moderate
holding capacity and high permeability, slightly acidic to hazards (Madulid and Agoo 1990). More specifically,
mildly alkaline and moderate to high fertile soils support available GIS proximity data analysis showed that on the
the Philippine teak forest (Ecosystems Research and average, the PTF grows inland at low altitude (142 m asl.)
Development Bureau 2003; Hernandez et al. 2016). All as far as 1.95 km from the coast; along steep ridges or hilly
these, together with other attributes of limestone substrates, to mountainous areas (52.37% slope) and very far
 CARINGAL et al. – Trees of Philippine teak forest 3197

(2.038km) from major riparian vegetation. The forest ACKNOWLEDGEMENTS

formations in the Philippines were classified according to
physiographic conditions from sea level up to the highest This study was made possible by the Accelerated
mountain (DENR-UNEP 1997; Malayang 2001; Pulhin 2002; Science and Technology Human Resource Development
Fernando et al. 2008). Richardson et al. (1995) stressed that Program – National Science Consortium (ASTHRDP-
soils and topographic conditions explained well the patterns of NSC) of the Department of Science and Technology
distribution and co-existence among plant species. (DOST) through the Graduate School and School of
Pausas and Austin (2001) suggested that studying Environmental Science and Management, University of the
multidimensional environmental factors can help Philippines Los Baňos (GS/SESAM, UPLB), Philippines.
understand the plant species composition in the landscape. The portion of botanical herbarium collections in Lobo,
This generalization was particularly important for the PTF Batangas was also covered by the research fund of
as other factors are affecting this threatened biota. In Batangas State University (BatStateU), Philippines.
particular, the forest patches are generally far (1.4 km)
from the settlement but near, 129.4m to cultivated hilly
lands and affected by road expansion 364 m away. Patches REFERENCES
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