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Journal of Asia-Pacific Biodiversity xxx (2018) 1e23
Contents lists available at ScienceDirect
Journal of Asia-Pacific Biodiversity
journal homepage: http://www.elsevier.com/locate/japb
Original Article
Plant diversity and structure of forest habitat types on Dinagat Island,
Philippines
Q22,2
Q3
Q4
Edgardo P. Lillo a, b, *, Edwino S. Fernando b, Mary Jane R. Lillo c
a
Cebu Technological University, Argao Campus, 6021, Cebu, Philippines
Forest Biological Sciences, College of Forestry and Natural Resources, University of the Philippines Los Baños, 4031, Laguna, Philippines
c
Argao Central Elementary School, 6021, Cebu, Philippines
b
a r t i c l e i n f o
a b s t r a c t
Article history:
Received 16 January 2018
Received in revised form
6 July 2018
Accepted 11 July 2018
Available online xxx
Logging, mining, and land conversion together threaten the whole Dinagat Island. The diversity and
structure of forest habitat types on Dinagat Island, Philippines, were determined as basis for conservation
and management. Identification of forest habitat types was based on habitat’s main physical characteristics. Six forest habitat types were identified, covered by 432 native plant species classified into 87
families and 203 genera, 9% or 40 plant species were endemic to Dinagat Island. Of the 432 species, 58%
recorded in lowland evergreen forest, 16% in upper montane, 15% in forest over limestone, 6% in lower
montane forest (LMF), 4% in mangrove forest, and 1% in beach forest, with average species diversity of
3.32. The number of threatened species was higher in the lowland evergreen forest, while species
richness was higher in the LMF. Endemism increases from the lowland evergreen forest to the montane
forest. Trees in lowland evergreen forest were bigger, taller, and larger in basal area as compared with
other habitat types. The most dominant species was Xanthostemon verdugonianus Náves ex Fern.-Vill
with importance value of 9.857%. The resource of native plant species occurring on ultramafic outcrops was an asset for the mineral resource industry for site rehabilitation and conservation.
Ó 2018 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing
Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).
Keywords:
Dinagat Island
Diversity
Forest habitat types
Species composition
Structures
Q5
Rationale
Q6
Dinagat Island is a northern extension of the northeastern
mountains of Mindanao, forming an island separate from the
remainder of Mindanao during the Pliocene epoch (Dicserson 1928;
Taylor 1934, Heaney 1986), when the Eurasian Plate and IndianAustralian Plate collided with the Philippine Sea Plate (Hamilton
1973). Dinagat Island is thought to have formed as a part of a
land bridge between Northern Mindanao and Eastern Visayas
(Leyte, Samar, and Bohol) in the late Pleistocene epoch (Leviton
1963; Heany 1986), facilitating the migration of species from
Mindanao to Eastern Visayas and vice versa.
Dinagat Island is the third largest Island in the Mindanao
biogeographic subregion located in the north of northeastern
Mindanao (Villanueva 2009; Figure 1). The island is considered one
* Corresponding author.
E-mail addresses: lillo_edgardo@yahoo.com (E.P. Lillo), edwinofernando@gmail.
com (E.S. Fernando), marylillo@yahoo.com (M.J.R. Lillo).
Peer review under responsibility of National Science Museum of Korea (NSMK) and
Korea National Arboretum (KNA).
of the areas in the Philippines characterized as ultramafic outcrops
together with Palawan, Samar, Zambales, Zamboanga, Mindoro,
and Sulu (Baker et al 1992; Balce et al 1976). The Dinagat Island is
rich in chromitite deposits similar to Zambales and Palawan (Yumul
1992; Yumul et al 2000; Zhou et al 2000). Alluvial platinum-group
minerals have been panned also in Dinagat Island together with
Samar (Franco et al 1993; Nakagawa and Franco 1995).
Logging, mining, and land conversion together threaten the
whole Dinagat Island (CI, DENR, Haribon 2006; DENR 2014;
Figure 2). The Philippine Biodiversity Conservation Priority-setting
Program spearheaded in 2001 by the Department of Environment
and Natural Resources in collaboration with nongovernment organization conservation groups identified Dinagat Island as an
extremely and highly critical terrestrial conservation priority.
Mining can affect biodiversity throughout the life cycle of a project,
both directly and indirectly. However, it cannot be denied that the
mining industry also plays significant role for the industrial and
technological development and the subsequent socioeconomic
progress around the world (ICMM 2010).
Dinagat Island has a unique faunal and floral composition, with
a high level of endemism (Heaney et al 1982; Ross and Lazell 1990;
https://doi.org/10.1016/j.japb.2018.07.003
pISSN2287-884X eISSN2287-9544/Ó 2018 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open
access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Figure 1. Location of study site in the Philippines and sampling site distribution on Dinagat Island.
Hämäläinen and Müller 1997). Mounts Kambinlio and Redondo,
located 10o 11’ 59" North (10.20o), 125o 34’ 59" East (125.58o), with
total area of 10,000 hectare lies at an elevation of 0e936 m (Bird
Life International 2017), on the northern part of the island,
together are regarded as part of Key Biodiversity Area (KBA) (No.
93) which covers four municipalities of Dinagat Island namely,
Loreto, Tubajon, Libjo, and Cagdianao. This Key Biodiversity Area
supports one critically endangered, four endangered, 13 vulnerable,
and 28 species with restricted range in the area (CI; DENR; Haribon
2006). One of the aforementioned critically endangered species is
Crateromys australis (Ambal et al 2012). In addition, two more
species of native tree, Gomphandra dinagatensis and Gomphandra
ultramafiterrestris (Schori and Utteridge 2012), have been recently
discovered from the island. Thriving also on Mount Kambinlio and
Mount Redondo are the Philippine flying lemur (Cynocephalus
volans) and the Philippine brown deer (Cervus marianna), which are
both considered endangered species. Tarsier (Tarsius syrichta) is
also abound in the said two mountains (DENR 2014).
Based on the study of Amoroso et al 2009 in Hamiguitan Range
an ultramafic mountain of the Southern Philippines, four vegetation
types are identified namely dipterocarp, montane, typical mossy,
and mossy-pygmy forests. The range shows a total of 878 species,
342 genera, and 136 families. Of the 878 species, 163 are endemic,
34 threatened, 33 rare, and 204 species with economic value.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Q20
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Figure 2. Proportion of areas covered by mining operation with approved MPSA in the
province of Dinagat Island (Source: DENR 2014).
In the study of Proctor et al in Mount Guiting-Guiting, an ultramafic mountain on Sibuyan Island, Romblon Province,
Philippines, it was found that the lower montane forest (LMF) is
high of species richness with up to at least 111 species of tree
(10 cm dbh) per 0.25-ha plot. At 770 m and 860 m, the Dipterocarpaceae accounted for 12.9% and 14.7% of the basal area,
respectively. There is a surprisingly high representation of the
Sapotaceae (25.9% of the basal area) at 1240 m. At 1540 m, the
Araucariaceae (Agathis sp.) dominated one plot (37.1% of the basal
area), and the Myrtaceae dominated the other (72.4%).
This study is considered unique because there was not much
work carried out to obtain the biological information of the ultramafic rocks (Castillo 2004), and of the seven provinces identified as
ultramafic outcrops by Baker et al 1992, only Palawan, Sibuyan, and
Romblon have been studied. So far, no ultramafic island has been
studied in terms of their floral composition and diversity. Knowledge on the exact plant species composition and diversity is an
important prerequisite to understand the structure and function in
ultramafic habitat types, biogeographical affinities, and their conservation and management (Jayatissa et al 2002; Wang et al 2003).
Sustainable use and management of natural resource is intimately
linked to ecology, as each management system interferes with the
forest structures and processes (Schmidt 1982).
The study aimed at assessing the diversity and structure of
forest habitat types on Dinagat Island, Philippines. Similarity and
dissimilarity among forest habitat types are important in understanding ecological variation.
Materials and methods
Study area
Q11
The study was conducted on Dinagat Island, Philippines
(Figure 1). Dinagat Island borders the province of Surigao del Norte
and Leyte. Dinagat Islands posted a total population of 126,803 with
an average annual population growth rate of 1.72%, total number of
household of 24,202, and average size per household of 4.99 (NSO
2010). The island can be reached through sea transport with 16
nautical miles west of Surigao City.
Dinagat Islands have an approximate land area of 80,212 has. or
802.12 sq. km. more or less including Hibuson Island and approximately 47 islets under the jurisdiction of the municipalities that
comprise the new province (RA 9355). Pedregosa-Hospodarsky
(2009) reported that based on NAMRIA (1988) Dinagat Island still
has 58% forest cover, but now the island has only 34% forest cover
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left and these remaining forests were mostly within the claimed
areas of several mining companies.
Geologically, the island was composed of recent alluvium
derived from MioceneePalaeogene rock. The Island lies between
the Philippine Fault and the Philippine Trench, and hosts the largest
layered chromite deposits in Leyte-Samar-Dinagat region (MGB
et al 1990). The whole province boasts of metallic and nonmetallic deposits, mostly chromite and gold with other byproducts in
appreciable amounts (Provincial Development Council 1993).
Dinagat Island has a soil pH that ranges from neutral to acidic Q12
(pH 4.8 to 6.9); average OM was 2.25%; phosphorus 1 to 37 ppm,
textural grade dominated by Dinagat clay loam (70%), 20% classified as Cabatohan loam, and the remaining 10% classified as
Bolinao clay (Haribon 2004). This type of soil was frequently noted
with pH higher in the subsoil than in the topsoil. This soil was
often associated with subsoils high in aluminum, low in calcium,
and pH very near or below the zero point of charge; such subsoil
will adsorb only trace amounts of calcium, magnesium, or potassium because of low negative charge density (Mekaru and Ueliara
1972).
Identification of forest habitat types
In this study, the identification of forest habitat types was
patterned from the study of Fernando et al (2008). The different
forest habitat types were grouped together according to the main
physical characteristics of their habitats. These forest habitat types
include: tropical lowland evergreen rain forest, tropical lower
montane rain forest, tropical upper montane rain forest, tropical
subalpine forest, forest over limestone (FOL), beach forest,
mangrove forest, peat swamp forest, fresh water swamp forest,
tropical semievergreen rain forest, and tropical moist deciduous
forest. Field reconnaissance and transect walks were conducted to
identify and describe the different forest habitat types.
Establishment of sampling plots
The establishment of plots was made in every study site
(Figure 3). A maximum of five plots and a minimum of three plots
were established for a site where the plots were laid at 300 m apart.
Plots were laid out at interval of 200 meters in elevation. For all
plots, the size used was 20 m 10 m. However, the actual location
of plots depended on the accessibility of the site. Generally, 20
meters is the longest distance that can be accurately surveyed in a
dense forest (Dallmeier 1992). Plants inside the quadrat were
identified and then counted accordingly. A 2 m 2 m subplot was
laid out inside the 20 m 10 m quadrat for the inventory of herbs,
vines, and seedlings.
Plant species identification and conservation status
Voucher specimens for every individual plant within the plot
were collected and tagged. The collected specimens were brought
to the metallophytes laboratory of the Forest Biological Science
Department, College of Forestry and Natural Resources, University
of the Philippines Los Banos Laguna (UPLB) for proper identification
after drying it from the oven. The identification of sample specimen
was done through manuals, Herbarium comparison, Co’s Digital
flora of the Philippines, online literature and also through the
expertise of Dr. Edwino S. Fernando. Conservation status of the
species in Dinagat Island was determined based on DENR (DAO
2017e11) and IUCN (IUCN 2016e3) classification.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Figure 3. Location of study site in the Philippines and sampling site distribution on different forest habitat types/formation of Dinagat Island (GIS generated map; Landsat 8; www.
Earthexplorer.usgs.ph; NAMRIA; Philippine GIS data).
Measurement of trees
The data for forest structure include diameter breast height
(DBH), tree heights, crown diameter and basal area. Native trees
with diameters of 1 cm and above were measured in terms of their
DBH, total height, and crown (height and width). The measurement
of DBH was done with the use of diameter tape for larger trees and
tree caliper for smaller trees. For the total height of the trees, the
measurement was made by the use of Abney hand level. For the
crown height and width, their measurements were done through
estimation. The tree height and DBH were categorized into
different classes such as 1e10 cm, 10e20 cm, 20e30 cm, 40e50 cm,
and 50 cm and above (Lulekal et al 2008). Basal area was calculated
by using the formula: BA ¼ 0.7854 (d)2, where d is diameter at
breast height in meter (DENR formula).
Species distribution and mapping
The location and distribution of sampling site was indicated in
the map, as well as the location of each species in each sample plot,
for forest structure and density characterization. The ground coordinates of each plot were determined by using GPS. The plot was
oriented in north-east direction to have an easy estimation on the
local coordinates of individual tree within plot. The local
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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coordinates of each individual tree within the plot was determined
by adding the X and Y distances to plot coordinates (Bantayan et al
2016). Elevation of each sample plots was determined using the
GPS.
Data analysis
Diversity of plant species
Native tree species diversity was computed and interpreted by
using the Shannon diversity index (H’). Shannon diversity index
was sensitive to areas with fragmented forest like in Dinagat Island.
However, Simpson and Brillouin indices were also used in the study
for comparison.
The Multivariate Statistical Package (MVSP) software was used
to compute the H’ of all sample plots by entering their respective
tree species and their corresponding density values as well as total
number of tree individuals in each sample plot. Compilation of H’
values of all sample plots provide valuable information especially in
explaining the relationship between H’ in relation with the absence
or presence of anthropogenic or natural stress factors. The same
MVSP software was used also to determine the corresponding
Shannon evenness index.
Species composition similarity index analysis (Jaccard’s index and
Sorensen’s index)
The similarity and dissimilarity among forest habitat types were
determined using the Jaccard and Sorensen index of similarity and
dissimilarity, based on presence and absence of species. There are
more than 20 binary similarity measures now in the literature
(Cheetham and Hazel 1969). Two of the most often used
similarity coefficients for binary data were Jaccard’s index and
Sorensen’s index. The Jaccard similarity coefficient refers to the
presenceeabsence matrix. The Jaccard’s similarity index formula:
ISj ¼ a/a þ b þ c; where a ¼ number of species in common between
the stands; b ¼ number of species unique to the first stand; c ¼
number of species unique to the second stand.
The Sorensen’s index was very similar to the Jaccard index and
was first used by Czekanowski in 1913 and discovered anew by
Sorensen (1948). The Sorensen’s similarity index gives greater
weight to matches in species composition between the two samples than mismatches. The Sorensen’s similarity index formula:
ISs ¼ 2a/2a þ b þ c, where a ¼ number of species in common between the stands; b ¼ number of species unique to the first stand;
c ¼ number of species unique to the second stand.
Cluster analysis and ordination
Cluster analysis was implemented using XLSTAT, Version
2016.02.28451, in the Microsoft Excel environment. The general
data analytical methods performed in the study were modified
from Andersen et al (2009) and Legendre et al (2008).
Table 1. The identified forest habitat types of Dinagat Island (Fernando et al 2008).
Soil water
Localities
Soil substrate
Elevation
Dry land
Inland
Ultramafic rocks
Lowlands (50e600 m)
Limestone
Montane (500e750 m)
Montane (750e929 m)
Lowland
Water table
Coastal
Salt water
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Plant species density, dominance, frequency, and importance value
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All recorded data were stored in a Microsoft Excel database and
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analyzed quantitatively using Microsoft Excel statistics. Vegetation
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analysis was done using the formula of density, relative density,
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dominance or basal area, relative dominance, frequency, relative
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frequency, and the importance value index. The ecological impor85
tance of each species in relation to the total forest community was
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calculated by summing its relative density, relative dominance, and
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relative frequency (Curtis and Macintosh, 1951). These provide a
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better index than density alone regarding the importance or
function of a species in its habitat. Alternatively the I.V. can also be Q13 89
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used instead of density alone in computing the plot’s Shannon di91
versity index and evenness index.
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Result and discussion
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Forest habitat types
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As patterned from the study of Fernando et al (2008), six forest
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habitat type are identified on Dinagat Island namely upper
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montane forest (UMF) (750e929 m), LMF (500e750 m), lowland
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evergreen rain forest (50e600 m), FOL, beach forest, and mangrove
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forest (e.g. Table 1). The lowland evergreen forest is classified into
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two subhabitat type based on the height of the species; the lowland
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tall forest (LF) covered by trees that attained a height of more than 5
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meters, and the lowland scrub forest covered by shrub-type trees
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that mostly attained a height of less than 5 meters.
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The upper montane forest, LMF, and lowland evergreen forest all
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have the same soil substrate, the ultrabasic rocks. The FOL is the
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forest habitat type with limestone substrate and mostly found in
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lowland. The beach forest is the forest habitat type in coastal areas.
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Then the mangrove forest is salt water habitat type (e.g. Table 1).
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The six forest habitat types appear to be altitudinal in sequence or
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compressed in Mount Kambinlio and Mount Redondo comparable
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to relatively more inland and larger mountains (Grubb 1971; 1977).
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This phenomenon is called as “Massenerhebung effect”. Compres115
sion of vegetation zones also observed on all major ultramafic
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mountains in Sabah, such as in Mount Meliau (1336 masl) and
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Mount Tawai (1273 masl), but the effect is most pronounced on
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Mount Tambuyukon (2579 masl) (Grubb and Whitmore 1966;
Proctor et al 1988; Bruijnzeel et al 1993; Ashton et al 2003).
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Forest habitat types
Height of the species
Forest type
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Lowland evergreen forest
Tall lowland forest (>5 m)
Tropical rain forest
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Lowland scrub forest (<5 m)
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Lower montane
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Upper montane
Forest over limestone
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Beach vegetation
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Mangrove forest
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Differences in native tree species composition between sites
were assessed with floristic dissimilarity matrices: (1) presence/
absence (PRAB) and (2) species abundance (ABU) data. Community
composition was computed across forest habitat types for all native
trees. Cluster analysis of native tree community and composition
was designed by Steinhaus dissimilarity matrix and Sorensen
dissimilarity matrix. Steinhaus dissimilarity matrix was computed
using the BrayeCurtis method, double standardized by species
maxima and site totals of log-transformed species abundance data
for each site. The Sorensen dissimilarity matrix was computed with
the same methods but using presence/absence data.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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The causes on the occurrence of this phenomenon involved
mean temperatures and cloud formation (Walker and Flenley
1979). However, according to Grubb and Whitmore (1966); Proctor et al (1988); Bruijnzeel et al (1993); and Ashton (2003), the main
causes for the compression of vegetation zones are related to the
lowering of the cloud base and the frequency of mist, as a result of
higher humidity and not by the increased adiabatic lapse rate. The
lowering of the cloud base in turn results in higher precipitation,
lower mean temperatures, less solar radiation, and slower
decomposition rates of organic matter in soils leading to a build-up
of humus and peat, acidification, and potential nutrient deficiencies
(particularly nitrogen) (Proctor et al 1988; Aiba and Kitayama
1999).
The identification of the six forest habitat types in Dinagat Island
indicates that the island has diverse forest ecosystem. Almost 60%
of the forest habitat types identified by Fernando et al (2008) in the
Philippines are found on Dinagat Island, and 47% of the forest
habitat types identified by Whitmore (1984) in Tropical Far East
Asia are also found on the island.
Species composition and description among forest habitat types
Upper montane forest
UMFs of the ultramafic mountain of Dinagat Island are comparable to Mossy forest of the Philippine forest formation of
Fernando et al (2008). In Dinagat Island, UMF is found on the
peak of the ultramafic mountain (Mount Redondo) with an area
of 527 hectares (e.g. Table 2), lies at an elevation of 750e
922 m (10.35686 N; 125.64806 E). The forest is covered by shrublike trees ranging from 1 to 3 meters in height. Local people call it
“bonsai forest”. This “bonsai forest” is confined to exposed slopes
of Mount Redondo. Shrub-like trees are the characteristics of
ultramafic habitats; however, due to its higher elevation, forest
classified as upper montane has conformed to the structure of the
upper montane of the tropical rainforest by Fernando et al
(2008).
The UMF habitat types are dominated by 81 species classified
into 34 families and 53 genera. Out of 81 species, 73 are native
trees, 15 shrubs, 4 herbs, and 2 vines/lianas (e.g. Table 3). The most
represented families are Rubiaceae, Arecaceae, Myrtaceae, Clusiaceae, Melastomataceae, Phyllanthaceae, and Thymelaeaceae. The
most represented genera are Syzygium, Phyllanthus, Vavaea, Lyptospermum, Elaeocarpus, and Psychotria.
The forests are dominated by the species of Leptospermum
amboinense Reinw. ex Blume, Scaevola micrantha C Presl., Gymnostoma rumphianum (Jungh. ex Vriese) L.A.S.Johnson, and
Dacrydium beccarii Parl. Despite the ecological importance of this
forest type, a large portion of it has been cleared for mineral
extraction.
Lower montane forest
In Dinagat Island, the LMF habitat type is located in the lower
elevation of the ultramafic mountain (Mount Redondo) with an
area of 693 hectares (e.g. Table 2). It occurs at an elevation that
ranges from 500 to 750 m (10.38228 N; 125.62225 E). The forest
habitat type is characterized with sparsely distributed and
irregularly shaped tree species with heights ranging from 5e10 m
and diameter ranging from 1e20 cm.
The forest habitat type is covered by 98 plant species classified
into 35 families and 57 genera. Out of the 98 species, 82 are native
trees, 4 shrubs, 5 herbs, and 7 vines/lianas (e.g. Table 3). The most
represented families are Rubiaceae, Arecaceae, Myrtaceae, Clusiaceae, Apocynaceae, Phyllanthaceae, Euphorbiaceae, Anacardiaceae,
and Moraceae. The most represented genera are Ficus, Syzygium,
Phyllanthus, and Psychotria. The forest habitat types are dominated
by the species of Weinmannia urdanetensis Elmer, Alstonia parvifolia
Merr, Canarium asperum Benth. in JD Hook., ssp. Asperum var.
asperum, Gymnostoma rumphianum (Jungh. ex Vriese) L.A.S.Johnson, Calophyllum blancoi Planch. & Triana, Terminalia darlingii Merr,
Ficus ampelas Burm, Syzygium sp, and Xanthostemon verdugonianus.
The characteristic and description of the forest conformed to the
lower montane of the tropical rain forest as described by Fernando
et al (2008).
Lowland evergreen forest
The lowland evergreen forest habitat type of the ultramafic
mountains of Dinagat Island is unique as compared to the typical
tropical rain forest formation of the Philippine Islands by Fernando
et al (2008). In Dinagat Island, lowland evergreen forest is classified
further into two subhabitat types based on the height of the species
of similar elevation. The first subhabitat type is the LF with a total
height of more than 5 meters, and the second subtype is lowland
scrub forest with a height of less than 5 meters.
The LF is found in Paragua forest (1134017.7 N; 779650.6 E) of
the Municipality of Libjo with an area of 6,265 hectares, Cuarinta
watershed of the Municipality of San Jose (1108880.0 N; 786387.0
E) with an area of 28 hectares, Basilisa forest of the Municipality of
Basilisa (1113569 N; 784791 E) occupied an area of 223 hectares,
Balitbiton Forest (1145548 N; 785642 E), Panamaon and Esperanza
watershed (1152045 N; 787262 E), and Mount Kambinlio (10.23
16.08 N; 125.38 56.93 E) all found in the Municipality of Loreto with
an area of 6,283 hectares, and Barangay Guerrero of the Municipality of Tubajon (1135198 N; 782333 E), with an area of 1,574
hectares (e.g. Table 2). The forest habitat types occur at an elevation
range from 50 to 600 m. The forests are covered by thick and dense
vegetation, with larger and taller trees.
The forests are covered by 350 plant species categorized into 70
families and 140 genera. Of the 350 plant species, 279 are native
trees, 32 are herbs, 17 are shrubs and 21 are vines/lianas
Table 2. Land coverage (ha) of different forest habitat types per Municipality of Dinagat Island (Google earth 2017).
Municipality
Basilisa
Cagdianao
Dinagat
Libjo
Loreto
San Jose
Tubajon
Beyond coastlines
Grand total
Forest habitat types of Dinagat Island
Lowland evergreen
forests
Forests over
limestone
Forests over ultramafic
rocks
Mangrove
forests
223
9,492
3,113
7,027
6,265
6,283
28
1,574
915
353
113
2,464
1,177
271
1,794
6,501
4,904
846
2,962
23,865
7,086
25,353
101
624
79
719
166
74
485
141
2388
Upper montane
forests
Lower montane
forests
527
693
527
970
Others
2,768
6,128
1,107
4,563
3,138
1,713
1,775
21,191
Grand
total
7,120
23,624
1,298
20,511
16,887
2,933
8,588
141
81,102
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Table 3. Life forms and composition of plant species in different forest habitat types.
Forest habitat types
Limestone forest
Lowland forest
Lower montane
Upper montane
Beach forest
Mangrove
Total
Location
Lake Bababu
Ferdinand
Santa Cruz, Tubajon
Subtotal
Paragua forest
Cuarenta watershed
Basilisa forest
Cambinliw forest
Esperanza
Panamaon
Tubajon (Guerrero)
Mount Kambinlio
Scrub forest
Subtotal
Lower elevation Mount Redondo
Bonsai forest, Mount Redondo
Subtotal
Subtotal
Life forms
Classification
Native trees
Shrub
Herb
Vines/liana
Families
Genus
Species
109
108
53
126
432
270
250
136
97
56
50
90
70
349
82
73
13
10
339
0
2
0
5
5
8
4
0
1
0
0
0
4
21
4
15
1
0
15
4
9
0
9
23
16
10
3
1
0
0
0
5
37
5
4
0
5
40
2
2
0
4
21
2
14
3
2
2
0
1
5
26
7
2
2
2
38
37
41
21
50
52
44
42
26
25
22
15
33
55
125
35
34
15
13
87
54
60
29
88
81
73
67
38
35
28
21
49
77
217
57
53
15
15
203
65
75
33
144
119
101
81
48
40
33
25
55
84
434
98
94
16
17
432
Species are shared from other study site.
(e.g.Table 3). The dominant families are Rubiaceae, Myrtaceae,
Apocynaceae, Arecaceae, Lauraceae, Moraceae, Dipterocarpaceae,
and Salicaceae. The dominant genera are Psychotria, Ixora, Shorea,
Syzygium, and Pinanga. Twenty-five percent (25%) of the species are
recorded from Paragua forest of the Municipality of Libjo. The
Basilisa forest, Cuarinta watershed, and Mount Kambinlio recorded
15% of the species, respectively.
The species of the family Dipterocarpaceae, Podocarpaceae, and
threatened species are recorded in the area. These species include
Shorea falciferoides Foxw. ssp. falciferoides, Shorea guiso (Blanco)
Blume, Shorea palosapis (Blanco) Merr., Shorea polysperma (Blanco)
Merr., Diospyros longiciliata Merr., Xanthostemon verdugonianus
Náves ex Fern.-Vill., Xanthostemon bracteatus Merr., Tristaniopsis sp,
Greeniopsis megalantha Merr, Eurycoma longifolia Jack, Mal. Kibatalia stenopetala Merr, Afzelia rhomboidea (Blanco) S.Vidal, Dacrydium beccarii Parl., and Falcatifolium gruezoi de Laub. The
characteristics and description of Dinagat Island lowland evergreen
forest conform to the lowland evergreen forest of the tropical
rainforest of the Philippines which includes the dipterocarp and the
mixed dipterocarp forests as described by Ashton (1997).
The lowland scrub forest is found in the Municipality of Cagdianao (1097024 N; 794810 E) and Municipality of San Jose
mountainous areas (1110476 N; 784649 E). The forest occurs at an
elevation ranges from 100 to 300 m, covered by less dense vegetation. The scrub forest are dominated by shrub to small trees with
a diameter at breast height ranges from 1 to 10 cm and height
ranges from 2 to 6 m. However, the forest is distinguishable from
lowland evergreen forest because of its low stature growth form
and uniform height. Scrub forest is also found in other municipalities; however, large areas and very prominent scrub vegetations
are found only in these two sites.
The forests recorded 84 plant species classified into 55 families
and 77 genera. The most represented families are Arecaceae,
Meliaceae, Myrtaceae, Phyllanthaceae, and Rubiaceae. The dominant genera are Phyllanthus, Psychotria, Medinilla, and Syzygium. Of
the 84 plant species, 70 are trees, five herbs, four shrubs, and five
vines/lianas (e.g Table 3). Sixty-six percent (66%) of the species are
recorded in Cagdianao ultramafic forest. The rest of the species
(34%) are recorded on the forest over ultramafic rocks of the Municipality of San Jose.
Species recorded common to the two ultramafic sites
includes Canarium asperum Benth. In JD Hook., ssp. asperum var.
asperum, Gymnostoma rumphianum (Jungh. ex Vriese) L.A.S.Johnson, Calophyllum blancoi Planch. & Triana, Garcinia sp., Dillenia sp.,
Diospyros littorea (R Br.) Kosterm., Fagraea gitingensis Elmer., Scaevola micrantha C Presl., Medinilla myrtiformis (Naudin) Triana,
Vavaea sp., Memecylon sp., Artocarpus multifidus Jarret, Decaspermum vitis-idaea Stapf., Leptospermum javanicum Blume., Syzygium sp., and Wikstroemia indica.
Scrub forests on Palawan are also dominated by 2e5 m tall trees
similar to Dinagat Island. The species composition of the forest is
also unique and composed of the heavy metal indicators Scaevola
micrantha, Brackenridgea palustris, and Exocarpus latifolius
(Podzorski 1985).
Most plants in ultramafic outcrops undergo morphological adaptations to minimize water requirements and water loss and
nickel and magnesium uptake for them to survive in drought
conditions (Brady et al 2005). Such adaptations include a generally
low stature, small crowned canopy, and other characteristics such
as glaucous leaves and sclerophyllous and microphyllous
morphologies.
Forest over limestone
In Dinagat Island, FOL habitat type are found in Lake Bababu
(10.07131 N; 125.51164 E) of the Municipality of Basilisa with an
area of 915 hectares, Ferdinand of the Municipality of Loreto
(10.38000 N; 125.55714 E) with an area of 1177 hectares, and Santa
Cruz of the Municipality of Tubajon (1141955 N; 784791 E) with an
area of 1,794 hectares (e.g Table 2). The forest habitat type lies at an
elevation ranging from 20 to 150 m. The forest is covered by less
dense vegetation, dominated by small size trees and few large trees.
The forest is also characterized by the occurrence of large-size
bedrocks with shallow soil and undecomposed organic matters.
Lake Bababu of the Municipality of Basilisa, Ferdinand of the
Municipality of Loreto, and Santa Cruz of the Municipality of
Tubajon are the locations that support this forest type. These sites
registered 144 plant species belonging to 50 families and 88 genera
(e.g Table 3). The most represented families (arranged in
descending order) are Moraceae, Rubiaceae, Euphorbiaceae, Clusiaceae, Anacardiaceae, Lauraceae, Apocynaceae, Phyllanthaceae,
Rutaceae, and Araliaceae. The dominant genera are Ficus, Psychotria, Timonius, Osmoxylon, Phyllanthus, and Buchanania.
Of the 144 species, 126 are native trees, nine herbs, five shrubs,
and four vines/lianas. Thirty-nine percent (39%) of the species are
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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recorded in Ferdinand FOL of the Municipality of Loreto. Thirty-six
percent (36%) are recorded in Lake Bababu site. The 25% of the
species are recorded in Santa Cruz of the Municipality of Tubajon.
The forest habitat type is dominated by Premna serratifolia L.,
Vitex parviflora Juss, Actinodaphne intermedia (Elmer) Ined, Phyllanthus ramosii Quisumb. & Merr., Mussaenda anisophylla Vidal,
Lunasia amara Blanco, and Leucosyke capitellata (Poir.) Wedd.
The characteristics and description of the forest conformed to
the Philippines FOL as described by Fernando et al (2008) and
Whitford (2011). Forest over limestone has a geological composition of mostly raised sedimentary and metamorphic rocks; a
considerable part of it being limestone (Audley-Charles et al 1979).
mangrove associates (Lugo and Snedekar 1974; FAO 2007). The
number of true mangrove species is equivalent to only 26% from the
total record of the Philippines with 39 species. (Fernando and
Pancho 1980). The result is equivalent in number to the study of
Cañizares and Seronay (2016) which recorded 10 mangrove species
in barangay Imelda of the Municipality of Tubajon, Dinagat
Island and equivalent only to 71% of the species recorded in
neighboring island as in Negros Island with 14 species (Calumpong
1994). The dominant species are Rhizophora apiculata Blume., Rhizophora mucronata Lam., Avicennia officinalis L., and Sonneratia
ovata.
Species composition of Dinagat Island
Beach forest
In Dinagat Island, beach forest habitat type is found in the
Municipality of Loreto and Dinagat. The forest habitat type appear
or form a narrow strip along the sandy beaches of the seacoast
above the upper tidal limits and bordered by roads in the opposite
side. This forest formation conformed to the beach forest formation
of the Philippine by Fernando et al (2008).
The beach forest has a total of 16 species, categorized into 15
families and 15 genera. Among the 16 species, 13 are classified as
trees, one shrub, and two vines/lianas (e.g. Table 3). The most
represented families are Fabaceae and Lecythidaceae. The most
represented genus is Barringtonia.
In the Municipality of Dinagat, beach forest has 10 species
classified into 10 families and 10 genera. The principal tree species
includes Talipariti tiliaceus L., Ipomoea pes-caprae (L.) Roth, Morinda
citrifolia L., Canavalia rosea (Sw.) DC., Scaevola taccada (Gaertn.)
Roxb., Barringtonia racemosa (L.) Blume ex DC, Terminalia catappa L.,
and Millettia pinnata (L.) Panigrahi in Panigrahi & Murti.
While in the Municipality of Loreto, 14 species are recorded in
the beach forest, categorized into 14 families and 13 genera. The
most dominant species are Ipomoea pes-caprae (L.) R. Br., Morinda
citrifolia L., Canavalia rosea (Sw.) DC, Cocos nucifera L., Millettia
pinnata (L.) Panigrahi in Panigrahi & Murti., Premna serratifolia L.,
and Hernandia nymphaeifolia (J.Presl) Kubitzki.
Mangrove forest
The mangrove forest in Dinagat Island is found in Llamera of the
Municipality of Libjo (10.13963 N; 125.54962 E) with an area of 719
hectares, Municipality of Dinagat (1105538 N; 786117 E) with an
area of 79 hectares, Municipality of Cagdianao (1099124 N; 790191
E) with an area of 624 hectares, and in the Municipality of Loreto
(0785694 N; 115174632 E) with an area of 166 hectares, and Basilisa
(1113569 N; 784791 E) with an area of 101 hectares (e.g. Table 2).
The mangrove forest habitat type is covered by less dense vegetation with uniform height and diameter sizes.
The forest is dominated by 17 plant species classified into 13
families and 15 genera from the 15 sampling plots. Out of the 17
total species, 10 species are classified as true mangrove and seven
species classified as associate mangrove species. Of the 17 total
species, 10 are classified as trees, five are herbs, and two are vines/
lianas (e.g.Table 3). The result is relatively similar to the true
mangrove species of North and Central America, which combined
to have 10 species. Indonesia (43) and Malaysia (41) have diverse
mangrove species than the Philippines (FAO 2007).
The most represented genus is Rhizophora, and the family is
Rhizophoraceae for true mangrove species and Orchidaceae for
associate species. The most represented genera for true mangrove
species are comparable to the study of Calumpong and Menez
(1996), published that the most dominant genera in the
Philippines are Rhizophora, Avicennia, Bruguiera, and Sonneratia.
True mangrove species are those species that grow in the
mangrove habitat only, while those not restricted to this habitat are
Dinagat Island with a total land area of 81,102 (e.g. Table 2),
covered by 432 native plant species classified into 87 families and
203 genera. Of the 432 native species, 61% or 263 of the plant
species are endemic to the Philippines and 30% or 130 species are
recorded or prehistorically originated from nearby tropical floral
region such as the countries found along the Pacific and Oceania,
Australia, West Malesia, East Malesia, Wallacea, and Continental
Asia, which are also considered as native species because of no
human interaction involved during their introduction in the site.
The remaining 9% or 40 plant species are endemic to Dinagat Island
and some could possibly be new species. This corresponds to 7.39%
of the total number of endemic plants in the Philippines.
Out of the 432 plant species, 79% (341) are classified as native
trees, 9% are both herbs and vines/lianas species, and only 3% are
shrubs. Of 341 native trees, 208 tree species are endemic to the
Philippines, and 31 are endemic to Dinagat Island. This corresponds
to 8% of the total number of endemic tree species in the Philippines.
Philippines has 3000 endemic trees (Philippines: Biodiversity e
Plants 2005).
Philippines is considered as home of 3557 indigenous or native
species and 26 native genera. Of these, 3200 species are angiosperms, six gymnosperms, and 351 are pteridophytes (ferns and
fern allies) (Amoroso et al 2006, 2009; Madulid 1991). The Hamiguitan Range Wildlife Sanctuary in the Province of Davao Oriental
in Mindanao Island has a total of 163 (36.69%) endemic species of
vascular plants, which corresponds to 5.09% of the total number of
endemic plants in the Philippines, lower percentage as compared to
Dinagat Island.
Endemic species are native species whose distributions are
confined only within the geographic area of reference. Thus native
species includes both endemic species and nonendemic indigenous
species whose natural geographic ranges extend beyond the
geographic area of reference. All species recorded in an area could
be classified as native species and alien species. Native species
could be endemic and nonendemic, while alien species are
wittingly and unwittingly introduced by humans to the geographic
point of reference (Coile 2002).
Native plant species has adapted and interacted through long
periods of time to local conditions (including weather, soil, water,
nematodes, fungi, bacteria, viruses, insects, mites, birds, reptiles,
mammals, fire, etc.) (Coile 2002).
Species diversity of forest habitat types
Upper montane forest
Based on Shannon diversity index estimation, the UMF has a
computed diversity value of 3.478, with a relative value of high
diversity (MacDonald 2003); whereas, from Simpson diversity index, the UMF has a diversity index of 0.952, with a relative value
very high (MacDonald 2003), and from Brillouin’s index, the UMF
has a diversity index of 3.87, with a relative value of high diversity
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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(MacDonald 2003). The three diversity index proved that the UMF
has high species diversity and evenness index of 0.961 or 96% of
species are common or shared among plots within the habitat types
(Figure 4).
Lower montane forest
Based on Shannon diversity index estimation, the LMF has a
computed diversity value of 4.081, with a relative value of very high
(MacDonald 2003) as well as on Simpson diversity index (0. 980)
with a relative value of very high. Brillouin’s index estimated a
diversity index of 3.545, with a relative value of high diversity
(MacDonald 2003). The three diversity index proved that the LMF
has a diversity value of high to very high species diversity, with an
evenness index of 0.827 or 83% of species are common or shared
among plots within habitat types (Figure 4).
Q15
Q16
Lowland evergreen forest
In lowland tall forest covering the forest areas in Paragua, three
watersheds (Cuarinta, Panamaon, and Esperanza), Balitbiton forest,
Tubajon, and Mount Kambinlio, whose species diversity value
estimation from Shannon index is 3.83, Simpson index is 0. 850,
and Brillouin’s index is 3.4. The result on the estimation of species
diversity by the three diversity indices significantly proving and
indicating for a similar species diversity result. The result implies
that the LF has a relative value of high species diversity (MacDonald
2003). Paragua forest, Basilisa forest, and Cuarinta watershed have
the highest species diversity value. Paragua forest is covered by 119
species categorized into 52 families and 81 genera (e.g. Table 3). The
high diversity in lowland evergreen forest conformed to the
description of lowland evergreen forest in the Philippines as
described by Fernando et al (2008).
For the distribution of species among LF, almost 95% are shared
with an average evenness index value of 0.947. The result reflects
the fact that almost 95% of the species are common from one LF to
another lowland forest.
However, for lowland scrub forest covering the areas of Cagdianao and San Jose, it has a computed diversity value of 3.46 for
Shannon index, 0.954 for Simpson index, and 3.469 for Brillouin
index. The resulting Shannon, Simpson, and Brillouin indices
confirm also the high species diversity on the lowland scrub forest
(MacDonald 2003). Cagdianao has the highest species diversity
value as compared to San Jose. Cagdianao is more diverse because it
9
is covered by 84 plant species classified into 55 families and 77
genera (e.g. Table 3). For the species distribution and abundance
among the two sites based on evenness index estimation, the
computed value is 0.873. The result also indicates that 87% of the
species distributed at scrub forests are shared among each other.
Forest over limestone
FOL covering the areas of Ferdinand of the Municipality of Loreto, Lake Bababu of the Municipality of Basilisa, and Santa Cruz of
the Municipality of Tubajon has a computed species diversity value
of 3.88 for Shannon index, 0.975 for Simpson index, and 3.2 for
Brillouin index (Figure 4). The result on the estimation of species
diversity by Shannon index and Brillouin index also signifies that
species diversities are also high in FOL. Likewise, Simpson index of
diversity also indicates a very-high species diversity. The three diversity indices prove that the species diversity of the FOL ranges
from high to very-high species diversity (MacDonald 2003). Among
the three forests over limestone site, Ferdinand of the Municipality
of Loreto and Lake Bababu of the Municipality of Basilisa have the
highest species diversity values. FOL of Ferdinand is considered as
more diverse because it is covered by 75 species categorized into 41
families and 60 genera (e.g. Table 3). In terms of species distribution
and abundance among the three sites based on evenness index
estimation, a computed value of 0.966 sort of confirms earlier diversity values. The results indicate that almost 97% of the species
among the FOL is shared with each other.
Beach forest
Based on Shannon, Simpson, and Brillouin diversity index
computation, the beach forest habitat type of Dinagat Island has a
diversity value of 1.45, 0.926, and 2.26, respectively (Figure 4). Both
Shannon and Brillouin diversity index computation indicated that
beach forest habitat type has low species diversity value
(MacDonald 2003). However, Simpson diversity index estimation
claimed that Dinagat Island beach forest has high species diversity.
The result further indicates that Dinagat Island beach forest
habitat type has low species diversity because the forest is only
covered by 16 species, 15 families, and 15 genera (e.g. Table 3). The
result is comparable to mangrove forest, both forest has low species
diversity (Figure 4). Based on the result of the evenness index of
diversity, both beach forest in the Municipality of Loreto and
Dinagat has an evenness index of 0.956 or almost 96% of the species
Figure 4. Species diversity value (Shannon, Simpson, Brillouin, and Evenness) among forest habitat types.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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in both sampling sites are similar to each other or common among
them (Figure 4).
Mangrove forest
Mangrove forest of Dinagat Island covers the area of the Municipality of Cagdianao, Dinagat, Basilisa, and Loreto as well as the
Gas Inlet river of Barangay Llamera of the Municipality of Libjo. The
species diversity estimation using Shannon, Simpson, and Brillouin
indices with values 1.44, 0.915, and 2.274, respectively (Figure 4),
indicates low species diversity for the mangrove (MacDonald
2003). Simpson index estimated value suggests a moderate species diversity value. Furthermore, among the five mangrove forests,
the one at the Municipality of Dinagat has the highest species diversity value compared with the other mangrove study sites. For
the species distribution and abundance based on evenness index
estimation, the computed value 0.941 indicates that 94% of the
species among the mangrove forests are common among them
(Figure 4).
Diversity of plant species on Dinagat Island
Based on Shannon diversity index estimation, Dinagat Island has
an average species diversity of 3.32 with a relative value of high
species diversity (MacDonald 2003). Whittaker et al., 1954 elaborated that an ultramafic outcrops has high level of plant
endemism and distinct vegetation composition and structure
compared to surrounding nonultramafic areas. Kruckeberg (1986)
and Wong (2011) emphasized that an adaptation to insular soils
(ultramafics) provides (genetic) isolation, which coupled with
strong edaphic and climatic stresses promotes evolutionary divergence and speciation.
Diversity is a community attribute related to stability, productivity, and trophic structure (McIntosh 1967; McNaughton 1977;
Tilman 1996), as well as migration (Wisheu and Keddy 1996;
Caley and Schluter 1997; Colwell and Lees 2000). An area with
high species diversity results to a more stable and productive
ecosystem.
Conservation status of native plant species
Conservation status of the species on Dinagat Island is deter- Q17
mined based on DENR and IUCN Classification (e.g Table 4). Using
the latest DENR classification under DAO 2017e11, there are five
species which have conservation statues declared as critically endangered, six endangered species, nine vulnerable species, and
three as other threatened species. There are also three nontree
species recorded in the site which are already threatened and
categorized as Endangered (Nepenthes alata Blanco, Nepenthes
truncata Macfarl, and Paphiopedilum ciliolare (Rchb.f.) Stein.
Using the IUCN classification, there are two species considered
as Critically Endangered A1cd ver 2.3, one species categorized as
Endangered B1þ2c ver 2.3, three species categorized as Vulnerable
A1cd ver 2.3, and one species categorized as Lower Risk/near
threatened ver 2.3. In this method of classification (IUCN), the
conservation status of some species does not coincide with those of
the DENR.version. In IUCN, Shorea falciferoides Foxw. ssp. falciferoides is considered as Critically endangered, but under the DENR
classification, they are still in Vulnerable category. Some species are
categorized as already threatened under the DENR classification,
but in IUCN record, the species are declared as not yet been
assessed. The contradiction between IUCN and DENR could be due
to the scale of work with the former at the global scale. This
contradiction may lessen the probability of categorizing and
updating the species during critical periods.
Similarity and dissimilarity among forest habitat types (Jaccard and
Sorensen)
Index of similarity and dissimilarity among forest habitat types
is analyzed by using the Jaccard and Sorensen indices (e.g. Table 4).
Table 4. Conservation status of native plants on Dinagat Island.
No.
Species
Endemicity
Conservation status
DAO 2017-11
IUCN 2016-3
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Greeniopsis euphlebia Merr.
Greeniopsis megalantha Merr.
Diospyros longiciliata Merr
Villaria acutifolia (Elmer) Merr.
Xanthostemon bracteatus Merr.
Eurycoma longifolia Jack, Mal. ssp. eglandulosa (Merr.) Noot.
Gomphandra ultramafiterrestris Schori
Kibatalia stenopetala Merr.
Madhuca lanceolata (Merr.) Merr.
Vitex parviflora Juss
Xanthostemon verdugonianus Náves ex Fern.-Vill.
Agathis philippinensis Warb.
Glenniea philippinensis (Radlk.) Leenh.
Pterocarpus indicus Willd. forma indicus
Sararanga philippinensis Merr.
Shorea falciferoides Foxw. ssp. falciferoides
Shorea polysperma (Blanco) Merr.
Sindora inermis Merr.
Terminalia surigaoensis Merr.
Vaccinium gitingense Elmer
Cinnamomum mercadoi Vidal
Elaeocarpus dinagatensis Merr.
Orania decipiens Becc.
Sonneratia ovata Backer
Nepenthes alata Blanco
Nepenthes bellii Kondo
Nepenthes merrilliana Macfarl.
Nepenthes truncata Macfarl
Paphiopedilum ciliolare (Rchb.f.) Stein
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Dinagat
Widespread
Philippines
Widespread
Widespread
Widespread
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Dinagat
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
Philippines
CR
CR
CR
CR
CR
EN
EN
EN
EN
EN
EN
VU
VU
VU
VU
VU
VU
VU
VU
VU
OTS
OTS
LC
LC
EN
EN
EN
EN
CR
Not assessed
Not assessed
Not assessed
Not assessed
Not assessed
Not assessed
Not assessed
Endangered B1þ2c ver 2.3
Not assessed
Vulnerable A1d ver 2.3
Vulnerable A1d ver 2.3
Not assessed
Not assessed
Not assessed
Not assessed
Critically Endangered A1cd ver 2.3
Not assessed
Vulnerable A1d ver 2.3
Not assessed
Not assessed
Not assessed
Not assessed
Lower Risk/Near Threatened ver 2.3
Lower Risk/Near Threatened ver 2.3
Lower Risk/Least Concern ver 2.3
Endangered 1þ2e ver 2.3
Not assessed
Not assessed
Endangered A2acdþ3cdþ4acd; B2ab (ii,iii,v); 1 ver 3.1
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Lowland scrub forest and LMF has the highest index of similarity
among the forest habitat types which ranges from 29e45% of its
species composition, or 45% of its species are shared to each other.
Lowland scrub forest and UMF has low index of similarity as
compared to other forest habitat types and ranges from 16e27%,
indicating for a very high index of dissimilarity, implying further
that these two forest habitat types almost have different species
composition.
The average similarity of species among forest habitat types
ranges from 23e37%, and the average dissimilarity among forest
habitat types ranges from 63e77% (e.g. Table 5). The results indicate
for a high dissimilarity of species composition among forest habitat
types. Dissimilarity of species composition among habitat types
also indicated uniqueness of species. The result further implies that
forest habitat types on Dinagat Island are distinct from each other.
Cluster analysis among forest habitat (Steinhaus and Sorensen
dissimilarity matrix)
Native tree community and composition based on species
abundance (Steinhaus dissimilarity matrix)
The plots sampled clustered into seven groups correspondingly,
as function to various species compositions and locations of forest
habitat types (Figure 5). Steinhaus dissimilarity matrix shows the
different forest habitat type as distinct from each other that
conform to the result of the study. The 18 plots from lowland forest
(LEF) composed of the plots of Cuarinta watershed, Basilisa forest,
Balitbiton forest, Panamaon and Esperanza watershed, and the
three plots from the forest of Mount Kambinlio proved to have
distinctive native tree species association hence grouping together
as one clump. Each lowland forest main group in turn divides into
subgroups and again into sub-subgroupings.
For example, the plots from Basilisa lowland forest split into
subgroups from subgroup with the plots of Cuarenta watershed.
The plots from Balitbiton forest likewise form subgroups consisting
of plots coming from Tubajon lowland forest, Esperanza, and Panamaon watershed as well as the plot from Mount Kambinlio (LDMR
p1, LDMRp2, and LDMRp3). The presence of the plots of Mount
Kambinlio of the Municipality of Loreto in the cluster of the lowland
forest represents a transition zone from lowland forest to UMF
(Bonsai forest).
The plots from Paragua lowland forest (LDLp1, LDLp2, and
LDLp3) form a separate cluster as distinct, having different species
composition, and abundance from other lowland evergreen forest.
In Mount Redondo, the plots from UMF composed of BUFp1,
BUFp2, BUFp3, BUFp4, and BUFp5 are also distinct in terms of
species composition and abundance as they form a separate cluster
from the other forest habitat types particularly the LMF. The plots
from LMF composed of LEMRp1, LEMRp2, and LEMRp3 are also
11
forming a separate cluster distinct from the other habitat types. The
formation of several cluster group that are distinct from each other
in terms of species composition and abundance within Mount
Redondo proved that the Mountain is composed of several forest
habitat type, all compressed together in a small mountain range.
These findings signify further that Mount Redondo has unique
vegetation type and need to be conserved to protect the diverse
biodiversity in the mountain.
The plots from FOL and beach forest both in Lake Bababu
(LSLBp1, LSLBp2, LSLBp3), Ferdinand (LSFp1, LSFp2 ans LSFp3) and
Santa Cruz (LDT) are also distinct in terms of species composition
and abundance as they are forming one cluster separate from other
forest habitat types. The plots from mangrove forest (LDMp1,
LDMp2, LDCMp1, and LDCMp2 and LDCMp3) are also distinct in
terms of their species composition and abundance as they form a
separate cluster from other forest habitat types.
The six plots from Cagdianao and San Jose lowland scrub forest
have distinct species composition as they are clustered into one
single group representing the scrub forest over ultramafic rocks
(FoUr) separate from the cluster of Mount Redondo. The cluster of
San Jose and Cagdianao signify for a difference in species composition between the two clusters based on its abundance, making
them distinct from each other (Figure 5).
Native tree community and composition based on absence/presence
data (Sorensen dissimilarity).
In the clustering analysis of native tree community and species
composition based on presence/absence data (Sorensen dissimilarity matrix), all the plots sampled are mainly divided into three
groups which correspond to three forest habitat types and locations
such as the lowland tall evergreen forest, lowland scrub FoUr, and
FOL (Figure 6). The 20 plots from LF covering the Cuarinta watershed, Basilisa forest, Balitbiton forest, Panamaon and Esperanza
watershed, lower elevation of Mount Redondo, Paragua forest, and
three plots from the forest of Mount Kambinlio proved to have
distinctive native tree formation and association as all clustered
into one single group. The plots from each lowland forest are
further clustered into subgroups. The plots from Paragua forest
(LDLp1, LDLp2, and LDLp3) together with the plots from Cuarinta
watershed (LDSCW p1, p2, and p3), as well as one plot from Basilisa
forest (LDBp3) constitute one subgroup. The three plots from LMF
(lower elevation of Mount Redondo) (LEMRp1, LEMRp2, LEMRp3)
together with the plots from Mount Kambinlio, Tubajon, Balitbiton,
and three plots from Basilisa forest form another subgroup. The
inclusion of the plots from Mount Kambinlio, which are classified as
forest over ultramafic rocks, into the group of LF represents a
transition zone from lowland forest to ultramafic forest.
The plots from mangrove forest together with the plots from
FOL in Lake Bababu, Ferdinand, and Tubajon appear to have a
Table 5. Similarity and dissimilarity of native species among forest habitat types (Jaccard and Sorensen index).
Plot
1
Forest habitat
Limestone forest
Native
species
126
126
2
Lowland forest
279
279
3
Forest over ultramafic
145
rocks
145
145
4
Upper montane
94
5
Lower montane
98
98
Average similarity and dissimilarity
Comparison between habitats
limestone forestelowland forest
Limestone forestelower montane
Lowland forestescrub forest
Lowland foresteupper montane
Scrub foresteforest over limestone
Scrub foresteupper montane
Scrub forestelower montane
Upper montaneeforest over limestone
Lower montaneelowland forest
Lower montaneeupper montane
Common
species
%Similarity
Jaccard index
Sorensen index
Jaccard index
%Dissimilarity
Sorensen index
111
69
139
94
78
42
87
42
94
69
22
26
25
21
22
16
29
17
20
27
23
35
42
40
34
37
27
45
29
35
42
37
78
74
75
79
78
84
71
83
79
73
77
65
58
60
66
63
73
55
71
65
58
63
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
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LDSCWsp1
LDSCWp3
LDBp4
LDBp1
LDBp2
LDSCW p2
LDSCW p1
LDSCW p3
LDBp3
LDMRp5
LDLMR p4
LDLMR p1
LDTp2
LDLMR p3
LDLMR p2
LDLCp2
LDLCp1
LDLCp3
LDLp3
LDLp2
LDLp1
BUfp2
BUfp1
BUfp3
BUfp5
BUfp4
LEMRp3
LEMRp2
LEMRp6
LEMRp1
LSFp1
LDTp1
LSFp3
LSFp2
LSLBp2
LSLBp1
LDM p1
LDCM p1
LDCp3
LDCp2
LDCp1
LDSp2
LDSp1
LEF
PELF
UMF
LMF
FOL
MF
FOUR
0
0.2
0.4
0.6
0.8
1
1.2
1.4
1.6
1.8
2
Figure 5. Dendrogram of all plots based on Steinhaus’ dissimilarity (species abundance data) which correspond to seven forest habitat types: lowland evergreen forest (LEF);
Paragua evergreen lowland forest (PELF); upper montane forest (UMF); lower montane forest (LMF); forest over limestone (FOL); mangrove forest (MF) and forest over ultramafic
rocks (FOUR).
distinct native tree composition as they clustered into one single
group. The inclusion of mangrove forest to the cluster of FOL represents another case of a transition zone.
The 10 plots from lowland scrub FoUr from Cagdianao, UMF
(Bonsai forest of Mount Redondo), and San Jose (FoUr), all have
distinct native tree species composition as they are clustered into
one single group representing the forest over ultramafic rocks
(Figure 6).
Comparison between Steinhaus and Sorensen Dissimilarity Index
The results of cluster analysis by Steinhaus dissimilarity matrix
and Sorensen dissimilarity matrix imply that each matrix has
different analysis. Steinhaus dissimilarity matrix identified the
seven forest habitat types that conform to the result of the study. In
fact, Steinhaus has separated the Paragua LF as one cluster distinct
from the other lowland tall evergreen forest habitat types
(Figure 5). In this type of matrix, it shows that Dinagat Island is
diverse in terms of forest habitat types as well as on biodiversity
species. The analysis of Steinhaus matrix conformed to the result of
the study that each forest habitat types are distinct from each other.
Furthermore, Steinhaus dissimilarity matrix is sensitive to the
detailed features of each forest habitat types in relation to the main
focus of the study.
While the Sorensen dissimilarity matrix identified only few
cluster groups, in this matrix, Sorensen has fused forest habitat
types and lumped them together as one cluster, similar to the UMF
and LMF that are lumped together with lowland scrub FoUr as one
distinct cluster (Figure 6). The special features of each forest habitat
types are not recognized in Sorensen dissimilarity matrix.
The results of the Jaccard and Sorensen index of similarity and
dissimilarity support the findings of Steinhaus dissimilarity matrix
that each forest habitat type is distinct from each other. (e.g
Table 5). Both Jaccard and Sorensen indicate for a high or significant
species differences among the forest habitat type. The result further
implies for high species diversity in Dinagat Island. Whitmore
(1984) also signifies that biodiversity occupies a distinctive physical habitat which is mostly sharply bounded, like for instance in
ultramafic outcrops environment.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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LDCp2
LDCp1
LDSp2
LDSp1
BUfp2
BUfp1
BUfp3
BUfp5
BUfp4
13
FoUr
LDSCWsp1
LDSCWp3
LDM p1
LDCM p1
LSFp2
LDTp1
LSFp1
LSLBp2
LSLBp1
LSFp3
LDBp4
LDBp1
LDBp2
LDLCp3
LDLCp2
LDLCp1
LDLMR p4
LDLMR p1
LDMRp5
LDTp2
LDLMR p3
LDLMR p2
LEMRp6
LEMRp1
LEMRp3
LEMRp2
LDSCW p3
LDSCW p1
LDSCW p2
LDBp3
LDLp3
LDLp2
LDLp1
FoL
LF
0
0.5
1
1.5
2
2.5
Sørensen Dissimilarity (Presence/ absence data)
Figure 6. Dendrogram of all plots based on Sorensen dissimilarity (species presence/absence data) which correspond to three forest ecosystem [forest over ultramafic rocks (FoUr);
forest over limestone (FOL); lowland forest (LF)]. Comparison between Steinhaus dissimilarity index and Sorensen dissimilarity index.
Relative frequency, density, dominance, and importance value
Based on computation, the most dominant species is Xanthostemon verdugonianus Náves ex Fern.-Vill. with an importance value
of 9.857% (e.g Table 6). The species is also the most dominant in
terms of diameter, richness, and density. Based on observation and
record, this species is considered as the biggest and largest tree
species in Dinagat Island, particularly in LF (Paragua forest, Balitbiton forest, and in different watershed areas). The tree species
recorded a diameter of 120 cm and total height of 25 m in Balitbiton
lowland evergreen forest. Wildlings and sapling of the species are
scattered in the forest floor of the LF and even along roads.
Glochidion album (Blanco) Boerl, the second most dominant
species with importance value of 6.715%. The species are recorded
in three forest habitat types (lowland evergreen forest, FOL, and in
ultramafic forest). The species is a small tree attaining only a
maximum diameter of 15 cm based on record but their frequency is
abundant in the area. Species wildlings and sapling are also scattered in the forest floor.
There are also other dominant native tree species that were
recorded in other forest habitat type (e.g Table 6), whose wildlings
and sapling are also dominant in the forest floor. The species
include Melicope triphylla (Lam.) Merr. (5.749%), Syzygium sp.
(5.142), Phyllanthus sp. (4.955%), Freycinetia sp. (4.668%), and
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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109
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111
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121
122
123
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128
129
130
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Table 6. Relative density, dominance, frequency, and important value of native trees.
No.
Species
Relative density
Relative dominance
(basal area)
Relative
frequency
Importance
value
1
2
3
4
5
6
7
8
9
10
Xanthostemon verdugonianus Náves ex Fern.-Vill.
Glochidion album (Blanco) Boerl.
Melicope triphylla (Lam.) Merr.
Syzygium sp.1
Phyllanthus sp.
Freycinetia sp.
Rinorea bengalensis (Wall.) Gagnep. in Humbert
Timonius sp.
Shorea polysperma (Blanco) Merr.
Canarium asperum Benth. in JD Hook., ssp. Asperumvar
Asperum
Garcinia sp.
Gardenia sp.
Severinia paniculata (Warb.) Swingle
Tristaniopsis sp.
Calophyllum blancoi Planch. & Triana
Leucosyke capitellata (Poir.) Wedd.
Vavaea sp.
Dillenia sp.
Podocarpus sp.
Syzygium sp.
Tetractomia tetrandra (Roxb.) Merr.
2.531
2.004
1.722
1.617
1.652
1.547
1.511
1.336
1.301
1.230
7.276
4.560
3.370
2.970
3.100
2.717
2.717
2.027
2.135
1.719
0.051
0.152
0.657
0.556
0.202
0.404
0.051
0.606
0.051
0.303
9.857
6.715
5.749
5.142
4.955
4.668
4.279
3.968
3.486
3.253
1.336
1.160
0.914
1.125
0.984
0.984
0.738
0.844
0.703
0.808
0.633
2.027
1.528
0.949
1.437
1.100
1.100
0.619
0.808
0.561
0.742
0.455
0.455
0.253
0.758
0.051
0.152
0.101
0.556
0.051
0.455
0.152
0.455
3.817
2.941
2.620
2.613
2.236
2.186
1.913
1.703
1.719
1.702
1.542
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Figure 7. Structure and density of the lowland tall forest of Paragua, Libjo. A, plot 1; B, plot 2; C, plot 3.
Figure 8. Structure and density of lowland and lower montane forest. A, lower montane forest; B, Balitbiton lowland tall forest; C, lowland tall forest of Cuarinta watershed.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Rinorea bengalensis (Wall.) Gagnep. in Humbert (4.279%), Timonius
sp. (3.968%), Garcinia sp. (3.817%), Shorea polysperma (Blanco) Merr.
(3.468%), Canarium asperum Benth. in JD Hook. ssp. Asperumvar.
Asperum (3.253%), Tristaniopsis sp. (2.613%), Calophyllum blancoi
Planch. & Triana, (2.236%), Leucosyke capitellata (Poir.)Wedd.
(2.186%), and Severinia paniculata (Warb.) Swingle (2.620%).
There are also nontree species that are dominant and found
scattered in the forest floor of the different forest habitat type. The
species are Dinochloa acutiflora (Munro) Soenarko, Dendrobium sp,
Nepenthes sp, Flagellaria indica L., and Dicranopteris linearis (Burm.
f.) Underw. Importance value is a quantity that measures the degree
of significance of tree species in a given forest community and is
derived from three variables, namely density, cover, and frequency
(pers.com.Baguinon).
Dinagat Island forest structure
Upper montane forest
The UMF (bonsai forest of Mount Redondo) has an elevation
ranges from 750 to 929 m. It is the highest peak in Dinagat Island.
The habitat type is characterized as mountainous in topography
with a vegetation cover estimated 70 to 90% and canopy cover
ranges from 10e40% (Figure 9). Ninety-four percent of the tree
species in the said forest showed diameters ranging from 1 to
10 cm, and 6% showed diameters ranging from 10 to 20 cm. The
forest has an average basal area of 53 m2/ha (e.g. Table 8). Most
trees reveal having height ranging from 1 to 2 m with average
crown diameter of 1 m.
The tree height is very insignificant from its diameter or stunted
growth. The trees at 10 cm diameter have a total height of less than
1 m. However, the tree species below 800 m elevation possess
10 cm diameter and attained a total height of more than 2 m and
crown diameter up to 3 m (e.g Table 8).
The result implies that plants tend to reduce its structure
(diameter and height) as an adaptation to low water holding capacity in higher elevation that causes water stress (Proctor 1999;
Brady et al 2005) as well as wind velocity which is stronger in
elevated areas.
Lower montane forest
The LMF (lower elevation of Mount Redondo) has elevation
ranges from 500 to 750 m. The habitat type is characterized as
mountainous in topography with vegetation and canopy cover
estimated 40 to 50% (Figure 8). Eighty-two percent of the tree
species in the said forest showed diameters ranging from 1 to
15
10 cm, 12% ranging from 10 to 20 cm, and 6% ranging from 30 to
40 cm. The forest has an average basal area of 44 m2/ha (e.g.
Table 8). Most trees reveal having height ranging from 6 to 15 m
with a crown diameter of 4 m. The forest structure (diameter,
height, and crown) can be comparable to lowland evergreen tall
forest. The mixture of different diameter classes and height result
to the formation of vertical layering in the canopy related to lowland evergreen tall forest.
Lowland evergreen forest
The LFs in Paragua, Basilisa, Tubajon, Balitbiton, and Mount
kambinlio and the watershed of Panamaon, Esperanza, and Cuarenta have topography ranging from rolling to mountainous. However, the LF of the Municipality of Tubajon is described only as
rolling. The elevation of the lowland evergreen forest ranges from
50 to 600 m above sea level. The forests of Paragua and Mount
Kambinlio have the highest elevation at the peak of 600 m. The
vegetation cover is 40e80% (Figures 7 and 8), covered by trees e.g.
diameter of 1e10 cm (74%), 10.1e20 cm (19%), 20.1e30 cm (5%), and
3% for trees with a diameter of 30 cm and above (e.g. Table 7). Those
with diameters 1e10 cm has total heights ranging from 3 to 7 m
with corresponding average basal area of 210 m2/ha, while trees
with diameter of 10.1 to 20 cm attained total height of 7 to 8 m with
average basal area of 188 m2/ha. Trees with 30 cm diameter has
total height of 12 m with average basal area of 120 m2/ha (e.g.
Table 7). The result reveals that as trees increase in diameter and
height, their number decline (Appendix 1). The decrease in number
of individuals as height increase implies dominance of small-sized
trees in the forest, reflecting a possible high rate of regeneration
(Bekele 1993; Senbeta and Denich 2006).
Trees with diameters of 1e10 cm, 10.1e20 cm, and greater than
20 cm tend to have crown diameters of 2 m, 3 m, and 4 m,
respectively. The result implies that as trees increase their diameter classes, the forest canopy cover tends to close. The lowland
forest of Paragua and Balitbiton and Cuarinta and Esperanza
watershed have all these diameter classes and therefore are expected to vary in terms of canopy cover corresponding to the
formation of vertical stratification or layering in their respective
canopies. Vertical stratification allows greater penetration of
sunlight down to the forest floor to the regeneration of smaller
plants other than trees and allows the germination of more seeds
deposited in the ground. Tesfaye Burju et al (2013) added that
diameter class distribution of tree species demonstrated various
patterns of population structure, implying different population
dynamics among species.
Figure 9. Structure and density in ultramafic rocks and upper montane forest. A, Municipality of San Jose; B, upper montane forest (Bonsai forest) plot 1; C, upper montane plot 2.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Table 7. Diameter classes and total basal area of lowland tall evergreen forest.
D. Classes
Lowland evergreen forest
1e10
10e20
20e30
30e40
40e50
50 and
above
Total
Basal
area
(m2/ha)
42
7
0
0
0
0
534
120
34
5
6
3
76
17
5
.7
.9
.4
210
188
120
31
59
95
49
702
100
703
Basilisa
forest
Balitbiton
forest
Cuarenta
watershed
Panamaon
watershed
Esperanza
watershed
Guerrero,
Tubajon
142
11
6
0
2
1
91
3
0
0
0
0
66
5
0
0
0
0
35
13
7
3
2
1
85
11
3
0
1
0
61
9
10
0
0
0
12
61
8
2
1
1
162
100
71
61
100
80
85
Mount Redondo
Total %
Lower
Upper montane
montane forest
1e10
41
10e20
6
20e30
0
30e40
3
40e50
0
50 and above 0
Total
50
%
Mt.
Kam.
Table 8. Diameter classes and total basal area in lower and upper montane forest.
Diameter
classes
Total
Paragua
forest
107
7
0
0
0
0
114
2
Basal area (m /ha)
Lower
Upper
Total
montane montane
148
13
0
3
0
0
166
89
9
0
2
0
0
100
16
9
0
19
0
0
44
42
11
0
0
0
0
53
58
20
0
19
0
0
97
Elevation of the area also affects forest structure. For example, in
Mount Kambinlio toward elevation of 600 m altitude, corresponding declines in vegetation and canopy cover as well as tree
diameters and heights were observed. Bruijnzeel et al (1993)
emphasized that plant or vegetations at higher elevation undergo
morphological adaptation to reduce transpiration. Elevated areas
have low water holding capacity that causes water stress to plants
hence the corresponding adaptation.
The LF is dominated by Xanthostemon verdugonianus Náves ex
Fern.-Vill., Melicope triphylla (Lam.) Merr., Glochidion album
(Blanco) Boerl., Shorea polysperma (Blanco) Merr., Tristaniopsis sp.,
Terminalia darlingii Merr., Areca species and Gymnostoma rumphianum (Jungh. ex Vriese) L.A.S.Johnson. In the forest floor, the
seedlings of these species also dominate together with undergrowth species such as Dinochloa acutiflora (Munro) Soenarko,
Dendrobium sp. and Nepenthes species.
While the lowland scrub forest has an elevation ranges from 100
to 300 m above sea level. The forest has a topography that is
mountainous with vegetation cover of 40e60%, and canopy cover
ranges from 20e40% (Figure 10). One hundred percent of the tree
species in the said forest showed diameters ranging from 1 to 10 cm
(e.g. Table 10). The forest has an average basal area of 50 m2/ha.
Most trees reveal having height ranging from 1 to 2 m with an
average crown diameter of 1 m. The result conformed to the
characteristics of plants in ultramafic outcrops in the Philippines as
described by Fernando et al (2008).
Forest over limestone
The FOL of Dinagat Island is on rolling to mountainous topography. The forest lies at an elevation of 103 m above sea level, with a
vegetation cover of 50 to 70%, dominated by trees with diameters
ranging from 1 to 10 cm (84%) and 16% at diameters above 10 cm.
The average basal area is 151 m2/ha (e.g Table 9: Figure 11).
Comparatively, the vegetation in limestone forest is larger and
taller than ultramafic forest but smaller compared to the lowland
forest. The total height of trees ranges from 7 to 12 m with an
average crown diameter of 3 to 6 m. Trees belonging to Ficus callosa
Willd., Ficus variegata, Caryota cumingii, and Vitex parviflora are
those that reach 19 cm DBH and are also the tallest in the limestone
Figure 10. Structure and density of the lowland scrub forest over ultramafic rocks of Cagdianao. A, plot 1; B, plot 2; C, plot 3.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Table 9. Diameter classes and total basal area in forest over limestone.
Diameter classes
(cm)
Forest over limestone
Ferdinand
Lake
Bababu
Tubajon
1e10
10e20
20e30
30e40
40e50
50 and above
Total
68
3
0
0
3
0
74
36
22
1
1
1
0
61
53
0
0
0
0
0
53
Total
%
Basal area
(m2/ha)
157
25
1
1
4
0
188
84
13
0.50
0.50
2
0
100
62
39.3
4
6.3
39.3
0
151
Table 10. Diameter classes and total basal area in lowland scrub forest.
Diameter
classes (cm)
Ultramafic forest
San Jose
Cagdianao
1e10
10.1e20
20.1e30
30.1e40
40.1e50
50 and above
Total
54
0
0
0
0
0
54
74
0
0
0
0
0
74
Total
%
Basal area
(m2/ha)
128
0
0
0
0
0
128
100
0
0
0
0
0
100
50
0
0
0
0
0
50
forest. The forest floor is dominated by the seedlings of Rinorea sp.,
Sterculia sp, Scolopia sp., Diospyros littorea, Timonius sp., and Dinochloa acutiflora (Munro) Soenarko.
Beach forest
The beach forest of Dinagat Island is feasible only in two sites
(Loreto and Dinagat). The forest habitat types are covered only by
few species with a vegetation cover of 20%. Few large principal trees
are growing in the second facies of the forest. The species of Terminalia catappa L., Barringtonia asiatica (L.) Kurz, Hernandia nymphaeifolia (J.Presl) Kubitzki, Barringtonia racemosa, Talipariti
tiliaceum, and Milletia pinnata are the dominant principal trees with
a diameter ranging from 15 to 25 cm, total height ranging from 7 to
10 m, and average crown diameter of 4 m. The forest has an average
basal area of 0.049 m2/ha.
Mangrove forest
The mangrove forest of Dinagat Island appears in patches in all
of the municipalities. However larger areas and more intact
17
mangrove forests exist in the municipalities of Libjo, Loreto, Basilisa, Dinagat, and Cagdianao. The five mangrove forests have estimated vegetation cover ranging from 30 to 40% (Figure 12). These
are dominated by trees with diameter breast height (dbh) ranging
from 1 to 10 cm (69%), and 31% of the trees have diameter greater
than 10 cm (e.g.Table 11) and average basal area of 47 m2/ha. The
total height of the tree ranges from 3 to 8 m with an average crown
diameter of 1 to 5 m. Larger tree species are recorded in the Municipality of Basilisa mangrove forest (e.g. Table 11). Other
mangrove areas possess similar sizes and structure. The mangrove
trees in Dinagat Island are smaller and shorter as compared to other
mangroves in the Philippine archipelago.
Rhizophora apiculata, Rhizophora mucronata, Sonneratia ovata,
and Avicennia officinalis are the species with larger diameter, basal
area, height, and dominance. The forest floor is dominated by the
seedlings of the same species. There are also three species of the
family Orchidaceae identified in the mangrove forest of the municipalities of Libjo, Dinagat, Basilisa, and Loreto. Epiphytes are
mainly habituated in a complex light atmosphere (Martin et al
2001). This indicated further that mangrove forests of Dinagat Island are dominated by smaller size trees, smaller basal area, open
canopy, and low species density.
Unfortunately, the structure of mangrove forest on Dinagat Island is smaller and shorter as compared to other islands in the
Philippines, particularly in Puerto Princesa, bay with an average
tree diameter of 104.5 cm, height of 15 m, and basal area of 438 m2/
ha (Dangan-Galon et al 2016).
Forest structure of Dinagat Island
The forest structure of different forest habitat types varies in
tree diameter, height, and basal area. The trees in lowland evergreen forest have larger diameter, taller trees, and larger basal area
coverage as compared to other forest habitat types. The lowland
forest has a tree diameter that ranges from 1 cm to 30 cm and above
with a height reaching to a maximum of 12 meters and total basal
area of 708 square meter/hectare (e.g. Table 7; Figures 7 and 8). The
limestone forest has also larger and taller trees second to lowland
forest (e.g. Table 9; Figure 11).
Tree structure of a forest served to give insight into its stand
density (Podong and Poolsiri, 2013). Tree structure will further
describe and give information on the vertical stratification of the
forest in each habitat types. This vertical stratification of the tree
canopy can affect the growth of young trees on the ground surface,
especially that of saplings and seedlings as well as on wildlife
Figure 11. Structure and density of forest over limestone. A, Ferdinand plot 1; B, Ferdinand; C, Lake Bababu of the Municipality of Basilisa; D, Santa Cruz Municipality of Tubajon.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Figure 12. Mangrove forest structure and density. A, mangrove forest of Cagdianao; B, mangrove forest of Llamera, Libjo; C, mangrove forest of Dinagat; D, mangrove forest of Basilisa.
Table 11. Diameter classes in mangrove forest of Dinagat Island.
Diameter classes (cm)
Mangrove forest
Llamera
Cagdianao
Dinagat
Loreto
Basilisa
1e10
10.1e20
20.1e30
30.1e40
40.1e50
50 and above
Total
23
12
3
0
0
0
38
23
3
2
0
0
0
28
25
5
3
0
0
0
33
30
8
3
0
0
0
41
25
10
3
4
0
0
42
species; as could be observed in lowland evergreen forest and FOL,
more number of seedlings and sapling growing in the forest floor,
result in high species diversity.
Kitayama (1991) emphasized that vegetation on ultramafic
outcrops as observed in Sabah is varied. Such variation in vegetation types in ultramafic rocks is likely to be the result of several
(synergistic) edaphic and other site factors, including soil chemistry, water stress induced by the substrate, erosion, exposure, and
elevation (Whitmore 1975). This phenomenon also happens in
Dinagat Island particularly in forest over ultramafic rocks and in
UMF. The reduction in growth structure of the trees serves as their
adaptation in order to survive at low nutrient availability and water
stress (Brady et al 2005). These growth adaptation has been termed
the “serpentine syndrome” (Jenny 1980; Rajakaruna and Baker
2004).
Total
%
Basal area (m2/ha)
126
38
14
4
0
0
182
69
21
8
2
0
0
100
49.5
60
49.5
25
0
0
184
Critically Endangered category (5), Endangered (6), Vulnerable
(9) species, and other threatened Species (3). This corresponds
to 2.2 % of the 984 threatened plant species in the Philippines.
4. Jaccard and Sorensen index of similarity as well as the cluster
analysis suggest that differences among forest habitat types
were high or significant.
5. The forest structure of different forest habitat varies in tree
diameter, height, and basal area. The trees in lowland evergreen
forest has larger diameter, taller trees, and larger basal area
coverage as compared to other habitat types particularly in UMF
and forest over ultramafic rocks.
Conflicts of interest
Q18
The authors declare that there is no conflicts of interest.
Conclusion
Uncited reference
1. Six forest habitat types were identified namely UMF, LMF,
lowland evergreen forest, FOL, beach forest, and mangrove forest. These forest types were covered by 432 native plant species,
classified into 87 families and 203 genera.
2. Based on Shannon diversity index estimation, Dinagat Island has
an average species diversity of 3.32 with a relative value of high
species diversity, while beach and mangrove forest has low
species diversity value.
3. From 432 species, 22 were threatened (DENR Administrative
Order-2017-11). This threatened species were categorized into
Q21
Baillie et al., 2000, Baselga, 2012, Coleman, 1971, Coronas, 1920,
Hall, 2002, Hall, 2012, Heany and Rabor, 1982, IUCN, 2011, JACCARD,
1908, Kruckeberg, 1984, O’Dell and Rajakaruna, 2011, Proctor and
Nagy, 1992, Proctor et al., 1999, Whitford, 1911, Zotz and Hietz, 2001.
Acknowledgment
The author would like to acknowledge Dr. Edwino S. Fernando
and Dr. Marilyn O. Quimado, his thesis advisers who were always
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
Q19
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there for him, the personnel of the Metallophytes Laboratory Tita
Mel, Tita Edith, Jay, Kat, Edu, and Irish for their support in the
conduct of the study particularly in the preservation of herbarium
specimens. He would also like to acknowledge Tita Mel Gibe for
helping in the germination of the seeds of native plants from
sampling sites, The Philippine Tropical Forest Conservation Foundation (PTFCF) for the financial budget as thesis grants, and the
personnel of the PENROeDENR, Dinagat Island for their support in
the processing of gratuitous permit and in the gathering of data.
The author acknowledges his brother and sister (Isang, Junjun, and
19
Mayette) for their help during the gathering of data in Dinagat Island, Jiro, for his help in the preparation of maps, colleagues Ritche,
Steve, and Archiebald for their encouragement and motivation, and
his wife and son (Mary Jane and CJ) for their moral and financial
support during the conduct of the study.
Appendix
Appendix 1. Forest structure of lowland evergreen forest (basal area, average diameter, height, and crown).
No
Scientific name
Average
HT (M)
Average crown
diameter (M)
Total
frequency
Average
DBH (cm)
BA/
TREE
(M2)
Yotal basal
area (M2)
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
Actinodaphne intermedia (Elmer) Ined.
Afzelia rhomboidea (Blanco) S.Vidal
Aglaia aherniana Perkins
Alchornea rugosa (Lour.) Muell.-Arg.
Alstonia macrophylla Wall. ex G.Don.
Alstonia parvifolia Merr.
Anacardiaceae 1
Apocynaceae
Apocynaceae
Apocynaceae
Archidendron clypearia (Jack) Nielsen var. sessiliflorum
Ardisia sp. 1
Areca caliso Becc. in Elmer
Areca costulata Becc.
Areca sp.
Arthrophyllum cenabrei Merr.
Artocarpus multifidus Jarret
Artocarpus sp.
Bikkia montoyae Mejillano, Santor & Alejandro
Brackenridgea fascicularis (Blanco) Fern. ssp.
mindanaensis (Merr.) Kanis,
Brackenridgea fascicularis (Blanco) Fern.-Villar ssp. fascicularis
Bridelia sp.
Buchanania heterophylla K.Schum.
Buchanania insignis Blume
Buchanania microphylla Engl. In DC
Callicarpa sp.
Calophyllum blancoi Planch. & Triana
Calophyllum cucullatum Merr.
Calophyllum sp.
Calophyllum sp.1
Calophyllum sp.2
Canarium asperum Benth. in JD Hook., ssp. asperum var. asperum
Canarium sp.
Celtis philippensis Blanco
Cerbera manghas L.
Cheilosa montana Blume
Cinnamomum mercadoi Vidal
Cinnamomum sp.
Commersonia bartramia (L.) Merr.
Cratoxylum sumatranum (Jack) Blume
Croton leiophyllus Muell.-Arg.
Dacrydium beccarii Parl.
Dasymaschalon clusiflorum Merr.
Decaspermum sp.
Dillenia philippinensis Rolfe
Dillenia sp.
Dillenia sp.1
Diospyros littorea (R Br.) Kosterm.
Diospyros longiciliata Merr.
Ehretia microphylla Lam.
Elaeocarpus sp.
Elaeocarpus sp.1
Euphorbiaceae
Eurycoma longifolia Jack Mal. ssp. eglandulosa (Merr.) Noot.
Fagraea sp.
Falcatifolium gruezoi de Laub.
3
6
8
8
6
7
7
7
4
6
4
3
3
7
5
6
8
7
3
4
2
2
3
3
3
2
3
2
1
3
1
1
2
5
3
3
2
1
1
2
3
15
16
24
18
2
37
4
6
40
2
16
66
3
9
6
19
16
2
5
11.00
15.00
14.00
12.00
15.00
20.00
5.00
2.50
10.00
1.00
6.00
15.00
15.00
10.00
13.00
10.00
12.00
1.00
6.00
6.00
0.00950
0.01767
0.01539
0.01131
0.01767
0.03142
0.00196
0.00049
0.00785
0.00008
0.00283
0.01767
0.01767
0.00785
0.01327
0.00785
0.01131
0.00008
0.00283
0.00283
0.14255
0.28274
0.36945
0.20358
0.03534
1.16239
0.00785
0.00295
0.31416
0.00016
0.04524
1.16632
0.05301
0.07069
0.07964
0.14923
0.18096
0.00016
0.01414
0.00565
4
4
6
6
7
6
3
8
5
7
5
5
7
12
6
5
4
8
7
7
4
5
3
8
4
9
8
7
5
6
3
8
6
5
7
6
2
2
2
2
3
3
2
2
2
2
2
2
2
4
2
2
2
3
3
4
1
2
1
2
2
4
3
3
2
2
2
3
1
2
1
2
2
10
6
17
9
27
116
11
21
12
7
7
12
2
10
4
4
10
20
10
1
2
2
15
4
31
79
11
40
24
1
19
4
14
15
17
10.00
6.00
10.00
10.00
10.00
12.00
9.00
10.00
7.00
7.00
7.00
12.00
6.00
8.00
8.00
8.00
15.00
9.00
12.00
5.00
7.00
1.00
10.00
6.00
17.00
15.00
16.00
5.00
10.00
5.00
15.00
6.00
10.00
10.00
10.00
12.00
0.00785
0.00283
0.00785
0.00785
0.00785
0.01131
0.00636
0.00785
0.00385
0.00385
0.00385
0.01131
0.00283
0.00503
0.00503
0.00503
0.01767
0.00636
0.01131
0.00196
0.00385
0.00008
0.00785
0.00283
0.02270
0.01767
0.02011
0.00196
0.00785
0.00196
0.01767
0.00283
0.00785
0.00785
0.00785
0.01131
0.07854
0.01696
0.13352
0.07069
0.21206
0.06786
0.72524
0.16493
0.04618
0.02694
0.02694
1.35717
0.13006
0.05027
0.02011
0.02011
0.17672
0.12723
0.11310
0.00196
0.00770
0.00016
0.82467
0.01131
0.70364
1.39605
0.22117
0.07854
0.18850
0.00196
0.33576
0.01131
0.10996
0.11781
0.13352
0.16965
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(continued on next page)
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
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Appendix 1 (continued )
No
Scientific name
Average
HT (M)
Average crown
diameter (M)
Total
frequency
Average
DBH (cm)
BA/
TREE
(M2)
Yotal basal
area (M2)
58
59
60
61
62
63
64
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
Fibraurea tinctoria Lour.
Ficus ampelas Burm.
Ficus callosa Willd.
Ficus nota (Blanco) Merr.
Ficus pseudopalma Blanco
Ficus sp.
Ficus sp.1
Flacourtia rukam Zoll. & Mor.
Flacourtia sp.
Flacourtia sp.
Garcinia sp.
Garcinia sp.1
Garcinia sp.3
Garcinia rubra Merr.
Gardenia sp.
Glochidion album (Blanco) Boerl.
Glochidion sp.
Glochidion sp.1
Glochidion sp.2
Gnetum gnemon L.
Gomphandra sp.
Gonocaryum cognatum Elmer
Greeniopsis megalantha Merr.
Greeniopsis sp.
Guioa sp.1
Guioa sp.2
Gymnostoma rumphianum
(Jungh. ex Vriese) L.A.S.Johnson
Heterospathe philippinensis Becc.
Heterospathe sp.
Homalanthus populneus (Geiseler) Pax
Leea indica (Burm.f.) Merr
Leea quadrifida Merr
Leptospermum javanicum Blume
Leucaena leucocephala (Lam.) de Wit
Leucosyke capitellata (Poir.) Wedd.
Lithocarpus sundaicus (Blume) Rehder
Litsea sp.
Lunasia amara Blanco
Macaranga tanarius (L.) Muell. Arg
Madhuca lanceolata (Merr.) Merr.
Magnolia liliifera Baill.
Meliaceae
Meliaceae 1
Melicope triphylla (Lam.) Merr.
Melodinus fusiformis Champ. ex Benth.
Memecylon sp.
Mitrephora williamsii C.B.Rob
Mussaenda anisophylla Vidal
Myristica sp.
Myrsine oblongibacca (Merr.) Pipoly
Myrtaceae
Myrtaceae 2
Neonauclea calycina (Bartl. Ex DC.) Merr.
Orania decipiens Becc.
Ormosia sp.
Ormosia surigaensis Merr.
Osmoxylon dinagatense (Merr.) Philipson
Osmoxylon eminens (Bull.) Philipson
Osmoxylon yatesii (Merr.) Philipson
Palaquim sp.
Pandanus apoensis Martelli, in Elmer
Pandanus sp.
Pandanus sp.1
Pandanus sp.2
Parartocarpus venenosa Becc.
Phyllanthus ramosii Quisumb. & Merr.
Phyllanthus securinegoides Merr.
Phyllanthus sp.
Phyllanthus sp.1
Phyllanthus sp.2
Phyllanthus sp.3
Phyllanthus sp.4
5
3
7
2
6
2
6
15
6
5
9
5
3
4
5
5
8
6
5
5
4
5
10
7
5
7
8
2
1
4
2
2
1
2
3
3
3
4
2
1
2
3
2
2
2
1
1
1
2
2
3
2
3
3
15
4
10
1
1
2
13
3
17
18
1
45
7
30
66
45
15
22
5
5
1
10
7
30
5
6
4
6.00
14.00
2.00
8.00
5.40
12.00
14.00
7.00
12.00
10.00
11.00
4.20
12.00
14.00
10.00
15.00
12.00
11.00
11.00
4.00
7.00
9.10
12.00
8.00
13.00
10.00
10.00
0.00283
0.01539
0.00031
0.00503
0.00229
0.01131
0.01539
0.00385
0.01131
0.00785
0.00950
0.00139
0.01131
0.01539
0.00785
0.01767
0.01131
0.00950
0.00950
0.00126
0.00385
0.00650
0.01131
0.00503
0.01327
0.00785
0.00785
0.01131
2.61695
0.00031
0.00503
0.00458
0.14703
0.18473
0.06542
0.20358
0.00785
0.42765
0.00970
0.33929
1.01599
0.35343
2.65073
0.24881
0.04752
0.04752
0.00126
0.03848
0.04553
0.33929
0.02513
0.07964
0.03142
0.39270
5
8
5
5
6
8
3
5
7
6
5
5
6
7
8
3
2
8
6
5
5
7
6
6
7
7
7
8
7
6
6
7
3
8
5
6
6
5
16
3
6
5
4
4
6
2
1
1
2
2
4
1
1
3
4
1
1
2
2
3
1
2
2
1
2
2
2
3
3
2
2
4
3
2
3
2
1
1
2
2
2
3
3
3
1
1
2
1
2
2
50
6
8
30
35
7
10
3
14
15
8
6
3
30
20
4
4
20
1
11
1
23
29
30
18
18
30
16
15
6
25
25
30
23
9
6
10
7
2
20
15
90
5
2
14
8.00
8.00
9.00
6.00
11.00
5.00
8.00
15.00
16.00
5.00
7.00
14.00
15.00
16.00
1.00
1.50
15.00
5.00
8.00
5.00
13.00
15.00
15.00
15.00
12.00
15.00
13.00
10.00
12.00
10.00
8.00
7.00
9.00
18.00
15.00
13.00
15.00
15.00
7.00
8.00
10.00
6.30
6.30
9.00
4.00
0.00503
0.00503
0.00636
0.00283
0.00950
0.00196
0.00503
0.01767
0.02011
0.00196
0.00385
0.01539
0.01767
0.02011
0.00008
0.00018
0.01767
0.00196
0.00503
0.00196
0.01327
0.01767
0.01767
0.01767
0.01131
0.01767
0.01327
0.00785
0.01131
0.00785
0.00503
0.00385
0.00636
0.02545
0.01767
0.01327
0.01767
0.01767
0.00385
0.00503
0.00785
0.00312
0.00312
0.00636
0.00126
0.03016
0.04021
0.19085
0.09896
0.06652
0.01964
0.01508
2.01455
0.30159
0.01571
0.02309
0.04618
0.53015
0.40212
0.00031
0.00071
2.12058
0.00196
0.05529
0.00196
0.30528
0.51247
0.53015
0.31809
0.20358
0.53015
0.21237
0.11781
0.06786
0.19635
0.12566
0.11545
0.14632
0.22902
0.10603
0.13273
0.12370
0.03534
0.07697
0.07540
0.70686
0.01559
0.00623
0.08906
0.00126
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Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
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Appendix 1 (continued )
No
Scientific name
Average
HT (M)
Average crown
diameter (M)
Total
frequency
Average
DBH (cm)
BA/
TREE
(M2)
Yotal basal
area (M2)
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
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168
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
Pinanga copelandii Becc.
Pinanga philippinensis Becc.
Pinanga sp.
Pinanga sp.1
Pittosporum moluccanum (Lam.) Miq
Podocarpus sp.
Polyscias cenabrei (Merr.) Lowry & G.M.Plunkett
Premna serratifolia L.
Psychotria scaberula Merr.
Psychotria sp.
Psychotria sp.
Psychotria sp.2
Psychotria sp.3
Psychotria sp.4
Pterocarpus indicus Willd. forma indicus
Radermachera gigantea (Blume) Miq.
Radermachera sp.
Rhodomyrtus surigaoensis Elmer.
Rinorea bengalensis (Wall.) Gagnep. in Humbert
Rubiaceae
Rubiaceae sp. 1
Sapotaceae 1
Saribus rotundifolius (Lam.) Blume
Scaevola sp
Schefflera sp.
Scolopia sp.
Semecarpus sp.
Severinia paniculata (Warb.) Swingle
Shorea falciferoides Foxw. ssp. falciferoides
Shorea guiso (Blanco) Blume.
Shorea palosapis (Blanco) Merr.
Shorea polysperma (Blanco) Merr.
Symplocos sp.
Syzygium sp.
Syzygium sp.1
Syzygium sp.2
Syzygium sp.3
Syzygium sp.4
Syzygium sp.5
Terminalia darlingii Merr.
Terminalia surigaensis Merr.
Tetractomia tetrandra (Roxb.) Merr.
Timonius lanceolatus Merr.
Timonius sp.
Timonius sp.1
Timonius sp.2
Trema orientalis (L.) Blume
Tristaniopsis sp.
Vaccinium epiphyticum Merr.
Vaccinium gitingense Elmer
Vavaea sp.
Vavaea sp. 1
Vavaea sp. 2
Vavaea sp. 3
Voacanga globosa (Blanco) Merr.
Xanthostemon bracteatus Merr.
Xanthostemon verdugonianus
5
9
8
9
7
7
5
8
4
6
5
5
4
4
5
5
9
10
5
6
5
8
4
2
3
4
5
7
5
9
7
8
10
9
5
7
6
7
7
5
8
6
5
10
4
5
6
6
8
8
2
6
3
3
3
10
12
2
3
2
2
2
3
2
1
2
0
1
2
2
1
1
2
2
4
0
2
1
2
2
1
0
1
2
3
2
3
2
2
3
2
3
2
2
2
2
2
3
3
2
3
2
2
2
3
2
3
1
2
1
1
2
5
5
20
5
13
45
5
17
20
5
28
1
6
26
4
6
36
1
49
3
1
40
11
5
7
5
1
2
25
26
86
50
10
22
20
26
70
28
40
91
6
10
16
10
35
4
11
26
24
33
80
1
20
2
4
10
3
14
120
10.00
10.00
15.00
11.00
11.00
10.00
6.00
10.00
4.00
3.80
8.00
6.00
6.00
10.00
3.00
12.00
14.00
5.00
7.00
9.00
10.00
3.50
8.00
5.00
8.00
5.00
11.00
7.00
15.00
12.00
10.00
20.00
11.00
11.00
15.00
11.00
10.00
12.00
13.00
12.00
13.00
7.00
8.19
5.00
8.00
6.00
10.00
15.00
10.00
5.00
12.00
5.00
5.00
16.00
7.00
20.00
40.00
0.00785
0.00785
0.01767
0.00950
0.00950
0.00785
0.00283
0.00785
0.00126
0.00113
0.00503
0.00283
0.00283
0.03142
0.00071
0.01131
0.01539
0.00196
0.00385
0.00636
0.00785
0.00096
0.00503
0.00196
0.00503
0.00196
0.00950
0.00385
0.01767
0.01131
0.00785
0.03142
0.00950
0.00950
0.01767
0.00950
0.00785
0.01131
0.01327
0.01131
0.01327
0.00385
0.00527
0.00196
0.00503
0.00283
0.00785
0.01767
0.00785
0.00196
0.01131
0.00196
0.00196
0.02011
0.00385
0.03142
0.12566
0.03927
0.10210
0.79522
0.04752
0.16156
0.15708
0.01414
0.21991
0.00126
0.00680
0.13069
0.01131
0.01696
1.13098
0.00071
0.55418
0.04618
0.00196
0.15394
0.06998
0.03927
0.00673
0.02513
0.00196
0.01005
0.04909
0.24709
0.33097
0.88358
0.11310
0.17279
0.62832
0.24709
0.66523
1.73181
1.04537
0.71471
0.06786
0.13273
0.18096
0.13273
0.13470
0.02107
0.21795
0.13069
0.06786
0.25918
1.41372
0.00785
0.03927
0.02262
0.00785
0.01964
0.06032
0.05388
0.03142
25.13280
References
Aiba S, Kityama K. 1999. Structure, composition and species diversity in an altitudesubstrate matrix of rain forest tree communities on Mount Kinabalu, Borneo.
Plant Ecology 140:139e157.
Ambal RGR, Duya MV, Cruz MA, Coroza OG, Vergara SG, De Silva N, Molinyawe N,
Tabaranza B. 2012. Key biodiversity areas in the Philippines: priorities for
conservation.
Amoroso VB, Obsioma LD, Arlalejo JB, Aspiras RA, Capili DP, JJA Polizon, Sumile EB.
2009. Inventory and conservation of endangered, endemic and Economically
important flora of Hamiguitan Range, southern Philippines. Blumea 54:71e76.
https://doi.org/10.3767/000651909X474113. www.ingentaconnect.com/content/
nhn/blumea.
Amoroso VB, Aspiras RA, JJA Polizon. 2006. Participatory inventory and distribution
of endangered, endemic and economically important plants in Hamiguitan
Range Wildlife Sanctuary, Davao Oriental. In: Proceedings of the annual in-house
review and evaluation of on-going and completed researches for the year 2006.
Musuan, Bukidnon, Philippines: University Research and Extension Office,
Central Mindanao University.
Andersen C, Nielsen TS, Purup S, Kristensen T, Eriksen J, Søegaard K, Sørensen J,
Frette XC. 2009. Phyto-oestrogens in herbage and milk from cows grazing
white clover, red clover, lucerne or chicory-rich pastures. Animal 3 (8):1189e
1195.
Ashton EC, Macintosh DJ, Hogarth PJ. 2003. A baseline study of the diversity
and community ecology of crab and molluscan macrofauna in the Sematan mangrove forest, Sarawak, Malaysia. Journal of Tropical Ecology 19:
127e142.
Ashton PS. 1997. Before the memory fades: some notes on the indigenous forests of
the Philippines. Sandakania 9:1e19.
Audley-Charles MG, Carter DJ, Barber AJ, Norvick MS, Tjokrosapoetro S. 1979.
Reinterpretation of the geology of Seram: implications for the Banda Arc and
northern Australia. Journal of the Geological Society 136:547e568.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
JAPB327_proof ■ 13 August 2018 ■ 22/23
22
EP Lillo et al. / Journal of Asia-Pacific Biodiversity xxx (2018) 1e23
Baillie IC, Evangelista PM, Inciong NB. 2000. Differentiation of upland soils on the
Palawan ophiolitic complex, Philippines. Catena 39:283e299.
Baker AJM, Proctor J, Van Balgooy MMJ, Reeves RD. 1992. Hyperaaccumulation of
nickel by the flora of the ultramafics of Palawan, Republic of the Philippines. In:
Baker AJM, JProctor, Reeves RD, editors. The vegetation of ultramafic (serpentine)
soils. Proceedings of the first international conference on serpentine ecology. U.K:
Intercept Ltd. Andover. pp. 291e304.
Balce GR, Alcantara PH, Morante EM, DH Almogela DH. 1976. Tectonic framework of
the Philippine archipelago (A review. Manila: Philippine Bureau of Mines Report.
59 p.
Bantayan CN, Cambalicer EA, Cltiburan JR, Barua LD, Dida JJV. 2016. GIS in the
Philippines. Principles and application in forestry and natural resources. 2nd ed.
Baselga A. 2012. The relationship between species replacement, dissimilarity
derived from nestedness, and nestedness. Journal of Macroecological Methods.
https://doi.org/10.1111/j.1466- 8238.2011.00756.
Bekele T. 1993. Vegetation ecology of renunant afromontane forests on the central
plateau of shewa, Ethiopia. PhD Dissertation. Uppssala, Sewden: Uppsala
University.
Bird Life International. 2017. Important bird areas factsheet: Mount Kambinlio and
Mount Redondo. Downloaded from, http://www.birdlife.org. on 20/04/2017.
Brady KU, Kruckeberg AR, Bradshaw Jr HD. 2005. Evolutionary ecology of plant
adaptation to serpentine soils. Annual Review of Ecology, Evolution, and Systematics 36:243e266.
Bruijnzeel LA, Waterloo MJ, Proctor J, Kuiters AT, Kotterink B. 1993. Hydrological
observations in montane rain forests on Gunung Silam, Sabah, Malaysia, with
special reference to the ‘Massenerhebung’ effect. Journal of Ecology 81:145e167.
Caley MJ, Schluter D. 1997. The relationship between local and regional diversity.
Ecology 78:70e80.
Calumpong HC, Menez EG. 1996. Field guide to the common mangroves,seagrasses
and algae of the Philippines. Makati City, Philippines: Bookmark Inc..
Cañizares LP, Seronay RA. 2016. Diversity and species composition of mangroves in
Barangay Imelda, Dinagat Island, Philippines. AACL Bioflux 9 (3):518e526.
Castillo ML. 2004. Uniqueness of vegetation in forest over ultramafic rocks and its relevance in forest restoration and rehabilitation or problematic areas in the Philippines.
http://agris.fao.org/agris-search/search.do?recordID¼PH2005000022.
CI, DENR- PAWB, Haribon. 2006. Priority sites for conservation in the Philippines: key
biodiversity areas; Quezon, city, Philippines conservation international Philippines.
Coile NC. 2002. Native plant? Wildflower? Endemic? Exotic? Invasive? Rare? Endangered? Botany Circular No. 35. Fla. Dept. Agriculture & Consumer Services. Division of Plant Industry.
Coleman RG. 1971. Petrologic and geophysical nature of serpentinites. Geological
Society of America Bulletin 82:897e918.
Colwell RK, Lees DC. 2000. The mid-domain effect: geometric constraints on the
geography of species richness. Trends in Ecology & Evolution 15:70e76.
Coronas J. 1920. The Climate and Weather of the Philippines, 1903 e 1918. Manila
Observatory. Bureau of Philippines.
Curtis JT, Macintosh RP. 1951. An upland forest continuum in the prairie forest
border region of Wisconsin. Ecology 32:476e498.
Dallmeier F, editor. 1992. Long-term monitoring of biological diversity in tropical
areas: methods for establishment and inventory of permanent plots. Paris: MAB
Digest. 11.UNESCO. 72 p. Retrievedfrom, http://unesdoc.unesco.Org/images/
0009/000938/093876eo.pdf. (Accessed 22 February 2008).
Dangan-Galon F, Dolorosa RG, Sespene JS, Mendoza NI. 2016. Diversity and structural complexity of mangrove forest along Puerto Princesa Bay, Palawan Island,
Philippines. Journal of Marine and Island Cultures. Journal of Marine and Island
Cultures 5 (2):118e125.
DENR Administrative Order. 2017. Updated national list of threatened philippine
plants and their categories (DAO No. 2017 -11).
DENR. 2014. DENR files, provincial office in Dinagat Island. Dinagat Island, CARAGA
region: PENRO.
Dickerson RE. 1928. Distribution of life in the Philippines. Philippine Bureau of
ScienceManila.
FAO. 2007. The World’s mangroves 1980e2005: a thematic study in the framework of
the global forest resources assessment 205. Rome, Italy: Food and Agriculture
Organization of the United Nations. pp. 1e74. Available online: http://www.fao.
org/docrep/010/a1427e/a1427e00.htm. (Accessed 25 February 2011).
Fernando ES, Suh MN, Lee J, Lee DK. 2008. Forest formation of the Philippines. ASEAN
e Korea Environmental Cooperation Unit (AKECU). GeoBook Publishing Co.
www.geobook.co.kr.
Fernando ES, Pancho JV. 1980. Mangrove trees of the Philippines. Sylvatrop, Philippine Forest Research Journal 5:35e54.
Franco HEA, Nakagawa M, Esguerra Jr E, Lazo FB, Domingo EG, Togashi Y. 1993.
Platinum-group minerals in laterite overlying the ultramafic massif in eastern
Samar and Dinagat Island, Philippines. Resource Geology 15:149e156. Sp l.
Issue.
Grubb PJ. 1971. Interpretation of the ‘Massenerhebung’ Effect on Tropical Mountains. Nature 229:44e45. https://doi.org/10.1038/229044a0. 01 January 1971.
Grubb PJ. 1977. Control of forest growth and distribution on wet tropical mountains,
with special reference to mineral nutrition. Annual Review of Ecological Systems
8:83e107.
Grubb PJ, Whitmore TC. 1966. A Comparison of Montane and Lowland Rain Forest in
Ecuador: II. The Climate and its Effects on the Distribution and Physiognomy of
the Forests. Journal of Ecology 54 (2):303.
Hall JL. 2002. Cellular mechanisms for heavy metal detoxification and tolerance.
Journal of Experimental Botany 53:1e11.
Hall R. 2012. SE Asia Research Group, Department of Geology, Royal Holloway University of London. Database files of geological bedrock supplied to Antony van der
Ent.
Hämäläinen M, Müller RA. 1997. Synopsis of the Philippine Odonata, with lists of
species recorded from forty islands. Odonatologica 26:249e315.
Hamilton W. 1973. Tectonics of the Indonesian Region. Bulletin of the Geological
Society of Malaysia 6:3e10.
Haribon. 2004. Foundation for the Conservation of Natural Resources, Inc. Dinagat
Technical Report: Mts. Kambinliw & Redondo, Loreto, Dinagat Island.
Heaney LR. 1986. Biogeography of mammals in SE Asia: estimates of rates of
colonization, extinction and speciation (1&2). In: Heaney LR, Patterson BD,
editors. Island biogeography of mammals. Biological Journal of the Linnean Society28. London, UK: Academic Press. pp. 127e165.
Heany LR, Rabor DS. 1982. Mammals of Dinagat and Siargao Islands, Philippines
Accepted for publication January 21, 1982. Occasional papers of the museum of
zoology University of Michigan.
Hospodarsky MP. 2009. Conservation of the Dinagat Tarsier (Tarsius syrichta carbonarius) and other threatened endemic mammals of Dinagat Island, Dinagat Province, Philippines.
ICMM. 2010. Integrating mining and biodiversity conservation: case studies from
around the World. Good Practice Guidance for Mining and Biodiversity.
IUCN. 2011. IUCN red list of threatened species. Version 2011.2.
IUCN. 2016. IUCN updates ’red list’ of endangered species. -3., https://www.iucn.nl/en/
solutions/red-list-of-threatened-species.
JACCARD. 1908. Nouvelles recherches sur la distribution florale. Bulletin de la Société
vaudoise des Sciences Naturelles 44:223e270.
Jayatissa LP, Dahdouh-Guebas F, Koedam N. 2002. A review of the floral composition and distribution of mangroves in Sri Lanka. Botanical Journal of the Linnean
Society 138:29e43. CrossRefWeb of Science Google Scholar.
Jenny H. 1980. The soil resource: origin and behavior. New York: Springer-Verlag.
Kitayama K. 1991. Vegetation of Mount Kinabalu Park, Sabah, Malaysia. Honolulu:
Environment and Policy Institute, East-West Center and Department of Botany,University of Hawaii at Manoa.
Kruckeberg AR. 1984. California serpentines: flora, vegetation, geology, soils and
management problems. Berkeley: University of California Press.
Kruckeberg AR. 1986. An essay: The stimulus of unusual geologies for plant
speciation. American Society of Plant Taxonomists 11 (3):55e463 https://doi.org/
10.2307/2419082. Stable URL: http://www.jstor.org/stable/2419082.
Legendre P, Borcard D, Peres-Neto PR. 2008. Analyzing or explaining beta diversity?
Comment. Ecology 89 (11):3238e3244.
Leviton AE. 1963. Remarks on the zoogeography of Philippine terrestrial snakes.
Proceedings of the California Academy of Sciences 42:112e145.
Lugo AE, Sneakers SC. 1974. The ecology of Mangroves. Annual Review of Ecology and
Systematics 5:39e64.
Lulekal E, Kelbessa E, Bekelet Yinger H. 2008. Plant species composition and
structure of the Mana Angetu moist montane forest, Southeastern Ethiopia.
Journal of East African Natural History 97:165e185.
Macdonald GM. 2003. Biogeography: space, time, and life. New York, NY: John Wiley
& Sons, Inc.
Madulid D. 1991. The endemic genera of flowering plants in the Philippines. Acta
Manilana 39:47e58.
Martin CE, Hsu R, Lin TC. 2001. Comparative photosynthetic capacity of abaxial and
adaxial leaf sides as related to exposure in two epiphytic ferns in a subtropical
rain forest in Northeastern Taiwan. American Fern Journal 99:145e154.
Mcintosh RI. 1967. An index of diversity and the relation of certain concepts to
diversity. Ecology 48:392e404.
Mcnaughton SJ. 1977. Diversity and stability of ecological communities: a comment
on the role of empiricism in ecology. American Naturalist 111:515e525.
Mekaru T, Uehara G. 1972. Anion adsorption in ferruginous tropical soils. SSSAP 36:
296e300.
Mines and Geosciences Bureau-Philippines (MGB) and Metal Mining Agency Of
Japan. 1990. Terminal report: The mineral exploration e minerals, deposits and
tectonics of two contrasting environments in the Republic of the Philippines.
Nakagawa M, Franco HEA. 1995. PGE abundance in ophiolitic rocks and soil from
Samar and Dinagat Islands,Philippines. GEOSEA 95 Pro. Sp l. Issue Journal of
Asian Earth Sciences.
NSO. 2010. Survey result. National Statistic Office.
O’Dell RE, Rajakaruna N. 2011. Intraspecific variation, adaptation, and evolution. In:
Harrison S, Rajakaruna N, editors. Serpentine: The evolution and ecology of a
model system. Berkeley and Los Angelis, California: University of California
Press.
Philippines: Biodiversity - Plants. 2005. Forest data: Philippines deforestation rates
and related forestry.CS. http://rainforests.mongabay.com/deforestation/archive/
Philippines.htm.
Podong C, Poolsiri R. 2013. Above ground biomass and litter productivity in relation
with carbon and nitrogen content in various landuse small watershed, Lower
Northern Thailand. Journal of Biodiversity and Environmental Sciences (JBES) 3
(8):121e132.
Podzorski A. 1985. The palawan botanical expedition: final report. 89p. Landskrona,
Sweden: Hilleshog Forestry AB.
Proctor J, Nagy L. 1992. Ultramafic rocks and their vegetation: an overview. In:
Baker AJ, Proctor J, Reeves RD, editors. The vegetation of ultramafic (serpentine)
soils. Andover, OR: Intercept. pp. 470e495.
Proctor J, Bruijnzeel LA, Baker AJM. 1999. What causes the vegetation types on
Mount Bloomfield? Global Ecology & Biogeography Letters 8:347e354.
Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
Asia-Pacific Biodiversity (2018), https://doi.org/10.1016/j.japb.2018.07.003
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
126
127
128
129
130
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
JAPB327_proof ■ 13 August 2018 ■ 23/23
EP Lillo et al. / Journal of Asia-Pacific Biodiversity xxx (2018) 1e23
Rajakaruna N, Baker AJM. 2004. Serpentine: a model habitat for botanical research
in Sri Lanka. Ceylon Journal of Science 32:1e19.
Ross CA, Lazell J. 1990. Amphibians and reptiles of Dinagat and Siargao islands,
Philippines. Philippine Journal of Science 119:257e286.
Schmidt P. 1982. Production ecology. In: Annual report, Project no. LH/UvS 01 Human
interference in the tropical rainforest ecosystem. Paramaribo, Suriname: CELOS.
pp. 30e43.
Senbeta F, Denich M. 2006. Effects of wild coffee management on species diversity
in the Afro montane rainforests of Ethiopia. Forest Ecology and Management
232:68e74.
Sorensen T. 1948. A method of establishing groups of equal amplitude in plant
sociology based on similarity of species content and its application to analyses
of the vegetation on Danish common. Kongelige Danske Videnskabernes Selskab
Biologiske Skrifter (Copenhagen) 5:1e34.
Taylor H. 1934. Philippine land mammals. Manila Bureau of Science, Department of
Agriculture and Commerce Monograph 30. 548 pp.
Tesfaye B, Hundera K, Kelbessa E. 2013. Floristic composition and structural analysis
of Jibat Humid Afromontane Forest, West Shewa Zone. Ethiopia: Oromia National
Regional State.
Tilman D. 1996. Biodiversity: population versus ecosystem stability. Ecology 77:
350e363.
Villanueva RJT. 2009. Adult odonata community in Dinagat Island, The Philippines:
Impact of chromium ore mining on density and species composition. Odonatologica 39 (2):119e126.
Walker D, Flenley JR. 1979. Late quaternary vegetational history of the Enga Province
of Upland Papua New Guinea, vol. 286. The Royal Society publishing. 1012.
23
Wang BS, Liang SC, Zhang WY. 2003. Mangrove flora of the world. Acta Botanica
Sinica 45:644e653. Google Scholar.
Whitford HN. 1911. The forests of the Philippines. Philippine Bureau of Forestry
Bulletin 10:113.
Whitmore TC. 1984. Tropical rain forests of the far east. 2nd edn. Oxford: Clarendon
Press. 352 pp.
Whitmore TC. 1975. Tropical rain forests of the far east. Oxford: Clarendon Press
Press.
Wisheu IC, Keddy P. 1996. Three competing models for predicting the size of species
pools: a test using eastern North American wetlands. Oikos 76:253e256.
Whittaker RN, Walker RB, Kruckeberg AR. 1954. The ecology of serpentine soils:a
symposium. Ecology 35:258e288.
Wong KM. 2011. A Revision Of Philippine Gardenia (Rubiaceae). Edinburgh Journal
Of Botany 68 (1):11e32. https://doi.org/10.1017/S0960428610000272 (2011)
11.Trustees of the Royal Botanic Garden Edinburgh 2011).
Yumul Jr GP. 1992. Ophiolite-hosted chromitite deposits as tectonic setting and
melting degree indicators: examples from the Zambales Ophiolite Complex,
Luzon, Philippines. Mining Geology 42:5e17.
Yumul Jr GP, Dimalanta CB, Jumawan FT. 2000. Geology of the southern Zambales
Ophiolite Complex, Philippines. The Island Arc 9:542e556.
Zotz G, Hietz P. 2001. The physiological ecology of vascular epiphytes: Current
knowledge, open questions. Journal of Experimental Botany 62:2067e2078.
Zhou MF, Yumul GP, Jr Malpas J, Sun M. 2000. A comparative study of platinumgroup elements in the Coto and Acoje blocks of the Zambales Ophiolite Complex, Philippines. The Island Arc 9:557e565.
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Please cite this article in press as: Lillo EP, et al., Plant diversity and structure of forest habitat types on Dinagat Island, Philippines, Journal of
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