taxonomic update on the begonia species of surigao …
TRANSCRIPT
TAXONOMIC UPDATE ON THE Begonia SPECIES OF SURIGAO DEL NORTE
Mico John Z. Dela Cruz
Mark Anthony Quintos
Submitted to the
Department of Biology
College of Arts and Sciences
University of the Philippines Manila
Padre Faura, Manila
In partial fulfillment of the requirements
For the degree of Bachelor of Science in Biology
June 2019
TABLE OF CONTENTS
List of Tables……………………………………………………………………………...……..iv
List of Figures………………………………………………………………………….…………v
List of Appendices……………………………………………………………………………….vi
Acknowledgment…………………………………………………………………………..……vii
Abstract………………………………………………………………….……………….…….viii
INTRODUCTION
Background of the Study……………………..………………........……………………...1
Statement of the Problem……………………………………………….…………………2
Research Objectives……………………………………………………………………….2
Significance of the Study……………………………………………………...…….….…2
Scope & Limitations of the Study……………………….…………………………….......2
REVIEW OF RELATED LITERATURE
Surigao del Norte and its location………….…………………...………………….…...4-5
Begonia…………………...………………………………...……………………...……6-7
METHODOLOGY
Preparation for the fieldwork and study sites………………………...……………….......8
Data collection procedure……………………..……………………………...…………8-9
Morphological analysis and data processing of the collected specimen…………........9-10
RESULTS
Results………………………………………………………………………………..11-32
DISCUSSION
Discussion……………………………………………….……………………………33-36
CONCLUSION & RECOMMENDATION
Conclusion and Recommendation……………………………………………………….37
REFERENCES………………………………………………………………...……………38-40
iv
List of Tables
Table
1 Major similarities and differences between Begonia acuminatissima (Rubite 999) and
B. acuminatissima Merr.……………………………...…………………………13-16
2 Major similarities and differences between Begonia colorata (Rubite 1000) and B.
colorata Warb………….……………………………………………………………..17-19
3 Major similarities and differences between Begonia sp. nov. 1 (Rubite 1001) and B.
obtusifolia Merr………….………………………………………………………...…20-23
4 Major similarities and differences between Begonia sp. nov. 2 (Rubite 1002) and B.
elmeri Merr………….…………………………………………………………….…25-27
5 Major similarities and differences between Begonia leucosticta (Rubite 1003) and B.
leucosticta Warb………….…………………………………………………………..29-31
v
List of Figures
Figure
1 Map of the Philippines…………………………………………………………………4
2 Map of Surigao del Norte………………………………………………………………5
3 Distribution of Begonia in Surigao del Norte………………………………………….12
4Begonia acuminatissima…………….…………………………………………………16
5 Begonia colorata …………………………….………..……………………………….20
6 Begonia sp. nov. 1……………………………………………………………………...24
7 Begonia sp. nov. 2 ………………………………………………………………..........28
8 Begonia leucosticta ……………………………………………………….…………...32
vi
List of Appendices
Appendix
A Wildlife Gratuitous permit of Mr. Mark Gregory Q. Rule………………………41-43
B Biographical Data………………………………………………………………..44-46
vii
Acknowledgements
We are thankful to God for the good health and well-being that He blessed us to complete
this undergraduate thesis.
We wish to express our deepest thanks to Mr. Mark Gregory Rule from University of the
Philippines Los Baños for joining and helping us during the botanical exploration.
We recognize the assistance of Cristoper M. Gales, Chairman of Barangay Motorpool,
and Peter Cortes who served as guides during our fieldwork.
We are grateful to the Department of Biology faculty for their help and support. We also
thank our parents, Mario and Ellen Dela Cruz, and Josephine Quintos for the moral, financial,
and emotional support to complete this research.
We are thankful to our panelists, Ma’am Fredeslinda C. Evangelista, PhD and Sir Jeffrey
P. Mancera, MSc, who provided us with further improvements to our study.
We would also like to express our gratitude for our thesis adviser, Rosario R. Rubite
Ph.D. for the unwavering supervision and guidance that were given to us.
viii
Abstract
Plant endemism in the province of Surigao del Norte has been poorly studied and no
taxonomic update with the current species it holds. Due to this, taxonomic claims of five species
of Begonia in Surigao namely Begonia bolsteri, B. leucosticta, B. longibractea, B. longistipula
and B. mindanaensis, were reinvestigated. This study revisited the area and conducted a study of
the genus Begonia, which included morphological characterization as well as preparation of
herbarium specimens. Five species of Begonia were collected: Begonia acuminatisima, B.
colorata, B. leucosticta, and two possible new species. Begonia sp. nov. 1 was compared with
another Philippine rugulose species, B. obtusifolia. Begonia sp. nov. 2 resembles another peltate
species found in Mindanao, B. elmeri. The new record of B. acuminatissima in Surigao del Norte
updates the distribution of Begonia species within Mindanao. The discovery of the presence of
these endemic begonias further promotes the diversity of species in the Philippines. The threats
that accompany with each species further reinforce the importance of their conservation.
1
Chapter I
INTRODUCTION
Background of the Study
The Philippines has one of the highest rates of wildlife endemism in the world. Out of the
52,177 recorded species, around 7,000 are endemic, which include 130 species of mammals, 190
species of birds, 196 species of reptiles, 95 species of amphibians, and around 6,200 species of
vascular flora (Philippine Species: A Unique and Distinct Multitude, 2014). However, these
endemic species are presented with a threat of extinction due to human activities. According to
the International Union for Conservation of Nature and Natural Resources (IUCN, 2018) Red
List, out of 785 threatened species, 245 of which are plants and the rest are threatened species of
birds, mammals, and amphibians. Illegal logging and mining provide the biggest threats through
destruction of the natural habitat of these native flora and fauna (Tacio, 2009)
Being the second largest island in the Philippines, Mindanao, particularly the Caraga
region, is recognized as a biodiversity hotspot that is currently threatened due to illegal logging.
(Farrell, 2016). The area of Surigao with its molave forests, ironwood forests, and mangrove
forests (Forest Management Bureau, 2016), boasts a number of flora and fauna endemic to it,
with the family of Begoniaceae being no exception. The five records of Begonia found in
Surigao are Begonia bolsteri Merr. (1911), B. leucosticta Warb. (Merrill,1911), B. longibractea
Merr. (1920), B. longistipula Merr. (1911) and B. mindanaensis Warb. (Merrill, 1911).
Presently, there have been no comprehensive studies on Begonia in the province. Being
residents near Surigao, it is fitting for the investigators to explore the area and study its Begonia
species.
2
Statement of the Problem
What is the current taxonomic status of the Begonia species in Surigao del Norte?
Research Objectives
The primary objective of the study is to make a taxonomic update of the Begonia species
in Surigao del Norte. Specifically, this study aimed to:
1. Collect Begonia species for the study
2. Record the morphological character of the collected species
3. Update the taxonomic description
4. Prepare a new set of herbarium collection
5. Investigate possible novel species during the botanical expedition
Significance of the Study
The study gives us an update on the present status of the Begonia species in Surigao del
Norte. It provides insights to the number of persisting Begonia species in their natural habitat.
The results confirm the species as recorded by Merrill (1911) and Warburg (1904). This also
provides an updated taxonomy of each species.
Scope & Limitations of the Study
This study was limited to taxonomic classification of the Begonia species in Surigao del
Norte based on morphological characters. The description of each Begonia species was done in
situ while other characters that cannot be measured or characterized on site were studied further
at the researcher’s field station. Collected and preserved specimens were submitted to the
Philippine National Herbarium and to the Department of Biology of University of the Philippines
Manila. Physiochemical tests, molecular analysis, and ecological studies were not included in
this study.
3
Chapter II
REVIEW OF RELATED LITERATURE
Surigao
The second largest of the three island groups in the Philippines, Mindanao, covers an area
of 97,530 km2 in total. Mindanao is primarily composed of mountain ranges, which include the
highest point in the Philippines, Mount Apo (Encyclopedia Britannica, 2016). Mindanao is also a
super region of the Philippines known as the “Agribusiness Mindanao”, which encompasses the
regions Zamboanga Peninsula, Northern Mindanao, Davao Region, SOCCSKSARGEN, Caraga,
and the Autonomous Region in Muslim Mindanao (ARMM) (Exec. Order No. 561, 2006). Most
of the land in Mindanao is used for agriculture, where the leading crops are corn, rice, abaca,
bananas, pineapples, mangoes, and coconut, and mining industry mostly owned by big
companies.
One of the regions in Mindanao, Caraga, is located on its northeastern portion. Caraga
covers a total of 18,846.97 km2 and comprises five provinces: Agusan del Norte, Agusan del Sur,
Surigao del Norte, Surigao del Sur, and Dinagat Islands and six cities: Butuan, Cabadbaran,
Surigao, Tandag, Bislig, and Bayugan. Caraga is also known for its agriculture and mining
industry, with their major products including rice, corn, coconut, banana, calamansi, gold,
silver,and iron. (Department of Trade and Industry, n.d.).
Surigao (Figure 1) was once a province of Caraga, which was split into two provinces,
namely, Surigao del Norte and Surigao del Sur (Republic Act 2786, 1960). Surigao combined
has a total area of 7,291.18km2 with Surigao del Norte having 37.5% of the total area (Figure 2)
and Surigao del Sur with 62.5%. The forests of Surigao cover a total area of 4,724.79 km2 and
can be considered as molave forests, ironwood forests, and mangrove forests (Forest
4
Management Bureau, 2016). Most of its forests are mossy watersheds and contain mineralized
soils, which are a suitable environment for the growth of Begonia (Island Properties n.d.). Its soil
types are primarily clay and sandy loam (Shin, n.d).
Figure 1. Map of the Philippines
Source:
https://earth.google.com/web/@9.79261156,125.80143509,35.34831894a,260185.54477505d,35y,0h,0t,0r/data=ChMaEQoJL20vMDFuazNsGAIgASgC
5
Figure 2. Map of Surigao del Norte
Source:
https://earth.google.com/web/@12.26182517,124.4642863,166.89086054a,2899181.78049535d,35y,0h,0t,0r/data=ChMaEQoJL20vMDFuazNsGAIgASgC
6
Begonia
The genus Begonia L. is popularly used worldwide as ornamental plant. It is distributed
globally in tropical and subtropical regions of the world (Naive, et al., 2018). Globally, 1,922
accepted species have been described and 70 sections been accepted but researchers believe that
many more can be discovered (Hughes, et al., 2015-). Asia, reaching as far from the Himalayas
to southern China and Malesia, is considered as a center for Begonia diversity, second to South
America. The last comprehensive account of Philippine Begonia by Merrill recorded 59 species,
but is now about a century old (Rubite & Madulid, 2009). As of 2018, a record of 132 species
(Hughes, et al., 2015-) has made the Philippines one of the centers of Begonia species diversity.
The Philippines provides the optimum conditions for Begonia species to grow, which include
plenty of indirect sunlight, humid environment, rich and aerated soil, and relatively cool weather
(Science Jrank, nd). Five species of Begonia that can be found in Surigao were identified. These
are B. bolsteri , B. leucosticta, B. longibractea, B. longistipula, and B. mindanaensis. Of the five,
B. longistipula is endemic to the area.
Begonia can be characterized as herbaceous plant that can either be a tall woody erect
shrub or a small fleshy rhizomatous herb, all of which with a fibrous root system. It has an
alternate phyllotaxy of asymmetrical leaves with a margin that can either be entire, toothed,
scalloped, or deeply jagged. It also has a cymose inflorescence that can either be axillary or
terminal and bracts that can either be large or small. The width of the monoecious flower ranges
at 1–5 cm, with the color of tepals ranging from pale pink to white. The male flowers have a
perianth composed of two outer tepals, opposite segments, and two inner smaller ones with
numerous stamens that are joined at the base, while the filaments are free or fused and the
anthers are narrowly obovate. On the other hand, female flowers usually have 5 tepals with the
7
outer tepals slightly larger than the inner ones. It also has an inferior ovary with two to three
styles placed above the tepals that are either free or fused and a stigma that can either be spiral or
twisted. Its fruit can either be fleshy or can be a dry winged capsule that is variously dehiscent or
irregularly breaking up. The seeds are very small and numerous (Kiew, 2005).
In the Philippines, there are currently two sections of Begonia, namely, Baryanda and
Petermannia. Species under section Baryandra are described as having thick rhizomatous stem
that are never erect. Their stipules are free and always persistent with all the leaves being
asymmetric and having palmate-pinnate venation, specifically actinodromous type. Their
inflorescences are axillary and cymose. They are protandrous, meaning the male flowers are
basal and open first while the female open last. Their flowers are always distal. Both the
staminate and pistillate flowers have two larger rounded outer tepals, oriented top and bottom;
and two inner tepals that are much narrower, and are oriented left and right (2+2), with their
filaments forming a short staminal tube. They have an inferior ovary that is 3-locular and the
style is divided into three branches that are each covered by a papillate, spiral, stigmatic band.
They have bilamellate placentas. Their fruits are 3-winged pendulous capsule and have either
equal or unequal wings (Rubite, 2012).
On the other hand, the section Petermannia is characterized as terrestrial, perennial plants
with erect stems, which are rarely rhizomatous. They have asymmetric leaves that are not peltate
and venation that ranges from palmate to pinnate. The inflorescence can either be axillary or
terminal that is usually bisexual wherein the male flowers are distal and the female flowers are
basal and often protogynous. The ovary typically has three equal or unequal wings that is often
3-locular, each containing placentas with two branches, and a pendulous capsule which are rarely
erect (Kiew, 2005).
8
Chapter III
METHODOLOGY
Fieldwork preparation
Ocular observations of the study site was made before the actual data collection. Prior
communication was done with Mr. Mark Gregory Rule of University of the Philippines Los
Baños who is currently doing a study entitled “Ecological and ethnobotanical characterization of
floral assemblages in forest over limestone formations of Surigao del Norte: Implications for
prioritizing local conservation strategies” in the area with a permit (Appendix A). Field materials
and equipment needed including Global Positioning System (GPS) device were prepared.
Descriptions of the known Begonia species from their protologues were studied prior to the
fieldwork.
Data collection
Data gathering was based on the study of Dastas & Guerrero (2018). Upon sight of a
Begonia, the exact coordinates and altitude of the collection sites were obtained using a GPS.
Descriptions of the habit, vegetative, and reproductive parts of the Begonia species were made
and photographs of every specimen in detail were taken in situ. Quantitative measurements of
each character were done in situ while further studies were done in the field station of the
researchers. Three flowering branches for each species were collected. Following Simpson
(2010), the collected specimens were laid between newspapers, applied with denatured alcohol,
sealed in airtight plastic bags and kept at room temperature. Upon arrival at the Department of
Biology of the University of the Philippines Manila, the collected specimens were placed in new
newspapers after which each specimen was dried by placing it between in two layers of
9
cardboard, pressed between two wooden frames that were tightly tied, and placed in an oven
with a temperature between 30-35°C. The collected herbarium specimens shall be submitted to
the Department of Biology and the Philippine National Herbarium.
Morphological analysis and data processing of the collected specimen
The morphological analysis was patterned from the study of Dastas & Guerrero (2018). A
12-inch Orion® ruler was used for the quantitative measurements of the vegetative parts of the
Begonia species. Measurement of each part when available was repeated three times and the
average was used. Measurements of the stem, leaf, and stipule were taken at the fifth internode
for uniformity. Without uprooting the plant, the stem length was measured from the base to the
apex. The diameter of the stem and internode length was recorded. The descriptions of the node
(e.g., presence of hair) were recorded. The dimensions of the stipules were measured and its
shape, color, presence of keel, apex, and presence of hair were noted. The length and diameter of
the petiole were measured and its color and presence of hair was recorded. The dimensions of the
leaf blade were measured from the base to its apex and widest width respectively. The blade’s
shape, apex, base, margin, and characteristics of its larger and smaller lobe were noted.
Descriptions of its adaxial and abaxial sides were also recorded.
The length and diameter of the peduncle were measured and its color and presence of hair
was noted. The bracts’ dimensions were measured and its shape and color was recorded.
Measurement of reproductive parts was observed for both the staminate and pistillate flowers.
The staminate flower’s dimensions were measured from the base to its apex and the widest
width, respectively. Its pedicel’s length and diameter were measured while the color and
presence of hair was noted. Measurement of the bracteoles’ dimensions and description of its
10
shape and color was recorded. The tepal’s number, shape, color, descriptions, of its adaxial and
abaxial side, was noted and its dimensions were measured. This is true for both the outer and
inner tepals. The union of the stamen, filament number, anther length, shape, filament length,
and height of stamen were recorded.
A parallel method was used for measurement of the parts of the pistillate flower.
Dimensions of the ovary were measured and its shape and color was noted. The presence of
wings and its description (e.g., shape, dimensions, color, apex, base, and presence of hair) was
recorded. The number of locules, types of placentation, length of style and color of the stigma
were recorded. The size of the fruit was measured by it dimensions and its shape, color, and
presence of hair was noted. The presence of wings and its description (e.g. shape, dimensions,
color, apex, base, and presence of hair) were recorded.
The data that were collected were cross-examined with the recorded catalogue of the
Begonia species described by Merrill (1911) and Warburg (1904). Taxon information of both
field and protologue data was compared and their differences noted. A more complete and recent
data is now produced.
11
Chapter IV
Results
Through the joint expedition with Mr. Mark Gregory Rule of the selected areas of
Surigao del Norte, a total of five Begonia species were found. With the aid of Google Earth, the
locations of the different species of Begonia were plotted to present the areas of the collected
species (Figure 3). These five Begonia species were identified through the use of the protologues
as described by Merrill (1911) and with the guidance of Dr. Rosario R. Rubite, the specialist on
Philippine begonias. All information regarding the vegetative characters were completed.
However, only two out of the five collected species had complete reproductive parts. Some of
the reproductive parts of the Begonia were still developing bud, while some do not have
reproductive parts during the fieldwork. Out of the collected Begonia species, two of these, B.
colorata Warb. (Merrill, 1911) and B. leucosticta, are native to Surigao Del Norte. The other two
are potential novel species, which are temporarily designated as Begonia sp. nov. 1 and Begonia
sp. nov. 2. While the last Begonia collected is a new record of B. acuminatissima Merr (1911).
The first Begonia (Rubite 999) was found in Barangay Motorpool, Municipality of Tubod
with numerous populations along the trail of a karst formation. Going up higher in the trail, the
second Begonia (Rubite 1000) was found but unlike the first Begonia seen, it was just a single
individual with only fruits present. The third Begonia (Rubite 1001) was discovered in Barangay
Sapa, Municipality of Claver, at the rock crevices of a riverine forest. The fourth Begonia
(Rubite 1002) was seen in Barangay Silop, Surigao City along the rock crevices at Silop Cave.
The fifth Begonia (Rubite 1003) was found in Barangay Sudlon, Municipality of Socorro at a
limestone formation near a coconut plantation and rice fields.
12
Figure 3. Distribution of Begonia in Surigao del Norte; Blue pin (Rubite 999), Red pin (Rubite 1000), Purple pin (Rubite 1001), Green pin (Rubite 1002), Yellow
pin (Rubite 1003)
Source: https://www.google.com/maps/d/edit?mid=1fyhm6SL_Kgs25nV9wFkXM14TTzelGevH&ll=9.668838812887559%2C125.69899065780032&z=11
13
` Begonia (Rubite 999) was observed to have a widely distributed population as numerous
clusters of individuals were found within the area of Barangay Motorpool. It was initially
identified as Begonia acuminatissima. It is a rhizomatous herb with narrowly ovate, prominently
acuminate leaves (Figures 4A & 4B), shiny glabrous adaxial side and pubescent abaxial side
(Figure 4B). Its petioles have dense hair (Figure 4C). The staminate and pistillate flowers were
also present (Figure 4D & 4E). Table 1 shows the major similarities and differences between the
data collected from the fieldwork and data from protologue of Merrill (1911).
Table 1. Major similarities and differences between Begonia (Rubite 999) and B. acuminatissima *
Collection Number Rubite 999 Merill 7306
Species name Begonia acuminatissima Begonia acuminatissima (Merrill, 1911)
STEM Rhizomatous Prostrate, creeping
Length 8 to 25 cm -
Diameter 7 mm -
Internode length 2 cm -
Color Brown -
Color Brown -
Hair Glabrous Glabrous
STIPULES - -
L x W 16 x 7.5 mm <1 cm L
Shape Triangular Oblong-ovate
Color Green Brown
Presence of keel Present -
Apex Aristate Acuminate
Hair Hair on margin -
Length 2 mm -
Color Maroon -
Distribution Dense -
LEAVES - Membranaceous
PETIOLE - -
Length 17.3 cm 8-18 cm
Diameter 5 mm -
Color Maroon -
Hair Present Few, ultimately glabrous
Length 0.4-0.5 -
Color Maroon Brown
Distribution Dense Scattered
BLADE - -
Shape Ovate Obliquely oblong-ovate or narrowly ovate
L x W 11.3 x 10.3 cm 7-11 x 3-6 cm
Apex Sharply acuminate Narrowed, long and sharply acuminate
Base Cordate Inequilaterally cordate
Margin Undulate Sinus is shallow, with small broadly triangular
lobes > 1cm in length
14
Larger lobe Undulate with small lobes Rounded
Smaller lobe Undulate Rounded
Adaxial Green color -
Hair Glabrous Glabrous
Abaxial Light green, shimmering -
Hair Present -
o Length 0.5 mm -
o Color Green -
o Distribution Sparse -
INFLORESCENCE - Dichotomously branched
PEDUNCLE - -
Length 20.7cm -
Diameter 3mm -
Color Green becoming reddish to the base -
Hair Present -
Length 1 - 1.5 mm -
Color Green -
Distribution Dense becoming sparse at the top -
BRACTS - -
Shape Ovate -
L x W 2.5 x 2.5 mm -
Color Green -
STAMINATE
FLOWER
- -
L x W 1.6 x 1.6 cm -
PEDICEL - -
Length 8mm -
Diameter 1mm -
Color Green -
Hair Glabrous -
BRACTEOLES Absent -
Shape - Broadly ovate
L x W - 1.5 x 1.5 mm
TEPALS - -
OUTER - -
Number 2 2
Shape Oblong Broadly elliptic, rounded
L x W 8 x 5 mm 7.5 x 6 mm
Color White Pink and white
Adaxial - -
Hair Glabrous -
Abaxial - -
Hair Glabrous -
INNER - -
Number - 2
Shape - Narrowly obovate
L x W - 6 x 4 mm
STAMEN - -
Union United at base -
Filament number Approximately 25 35
Anther length ~1 mm 1 mm
Shape Rounded Truncate
Filament length 1.5 mm 0.5-1 mm
15
Height of stamen 2.5 mm -
PISTILLATE
FLOWER
- -
L x W 1.6 x 1.6 cm -
PEDICEL - -
Length 1 cm -
Diameter 1 mm -
Color Green -
Hair Glabrous -
BRACTEOLES Absent -
TEPALS - -
OUTER - -
Number 2 -
Shape Oblong -
L x W 8 mm x 5 mm -
Color White -
Adaxial - -
Hair Glabrous -
Abaxial - -
Hair Glabrous -
INNER - -
Number 2 -
Shape Oblangulate -
L x W 8 mm x 3 mm -
Color White Pink and white
Adaxial - -
Hair Glabrous -
Abaxial - -
Hair Glabrous -
OVARY - -
Shape Oblong -
L x W 9 mm x 2.5 mm -
Color Green -
Wings Sub-equal wings -
Shape Rounded -
L x W 8 mm x 5 mm -
Color Green -
Apex Rounded -
Base Rounded -
Hair Glabrous -
Number of locules 3 -
Type of placentation Axillary 2-partite
Style Absent -
Stigma Absent -
FRUIT - -
Shape Oblong
L X W 9 mm x 2.5 mm 8-10 x 15 mm
Color Brown -
Hair Glabrous -
Wings Sub-equal Sub-equal
Shape Rounded Somewhat narrowed and subacute or blunt
L X W 8 mm x 5 mm 5-6 mm W
Color Green -
16
Apex Rounded Acute or blunt
Base Rounded Retuse
Hair Glabrous -
*green highlight = similarities, yellow highlight = updated characters, dash (“-“) = not seen or not stated in
protologue.
Figure 4. Begonia acuminatissima A. Habitat; B. Adaxial and Abaxial side; C. Petiole with dense hair; D. Staminate flower;
E. Pistillate flower
17
Begonia (Rubite 1000) initially identified as Begonia bolsteri. It was a lone, upright
Begonia species (Figure 5A) without staminate and pistillate flowers and only its fruits were seen
(Figure 5B). Table 2 shows the major similarities and differences between the data collected
from the fieldwork and data from Naïve et al. (2018), declaring B. bolsteri as synonym of B.
colorata.
Table 2. Major similarities and differences between Begonia (Rubite 1000) and B. colorata *
Collection Number Rubite 1000 Warburg 14633
Species name Begonia bolsteri Begonia colorata (Warburg, 1904)
STEM Erect Unbranched, erect at the apex, lower
point repent
Length 20 cm 15-30 cm
Diameter 0.4 cm 0.3-0.5 cm
Internode length 3.5 cm -
Color Purple Maroon
Node A little bit swollen Rooting
Color Purple to brownish
Hair Glabrous Glabrous
Length - -
Color - -
Distribution - -
STIPULES - Membranaceous
L x W 8 mm x 4 mm 0.7-1 cm L
Shape Triangular Oblong-ovate
Color Purple red Reddish brown
Presence of keel Present
Apex Acuminate Acuminate
Hair Glabrous
LEAVES - Alternate, succulent, terete, palmate-
pinnate
PETIOLE - -
Length 5.27 cm 6-13 cm
Diameter 2.7 mm
Color Purple-red red
Hair Glabrous
BLADE - Inequilateral, slightly membranaceous
Shape Obliquely ovate Oblong-ovate to narrowly ovate
L x W 8.57 x 5.33 cm 5.5-8 cm x 2-5 cm
Apex Acuminate Attenuate to acuminate
Base Cordate Obliquely cordate
Margin Angulate Irregularly and coarsely dentate
Larger lobe Small lobes at the tip of the veins/ Rounded
18
denticulate tips
Smaller lobe Denticulate tips Rounded
Adaxial Shiny. Dark green, but longest primary
vein is light green
Hair - ciliate
o Length - -
o Color
- Dull green with distinct pale grey green
stripes at the midrib, sometimes having
a dark mottled pattern
o Distribution - -
Abaxial Maroon
INFLORESCENCE - Terminal or axillary
PEDUNCLE - -
Length - 3-4.5 cm
Diameter - 1.5- 2cm
Color - Maroon
BRACTS - Membranaceous
Shape - Lanceolate
L x W - 0.5-0.8 x 0.2-0.3 cm
STAMINATE FLOWER - distal
PEDICEL - 5-8 mm
Hair - glabrous
BRACTEOLES - -
TEPALS - -
OUTER - -
Number - 2
Shape - Orbicular to reniform
L x W - 4.5-5 x 5-6 mm
Color - pink
STAMEN - -
Union - -
Filament number - 20-25
Anther length - 0.8-1 mm
Shape - Narrowly obovoid
Filament length - 1-1.5 mm
Height of stamen - -
PISTILLATE FLOWER - basal
L x W - -
PEDICEL - -
Length - 5-10 cm
Hair - glabrous
BRACTEOLES - -
TEPALS - 5
OUTER - -
Shape - Broadly ovate
L x W - 7-8 x 5 mm
Color - pink
19
INNER - -
Number - -
Shape - Broadly ovate
L x W - 7-8 x 5 mm
Color - pink
OVARY - -
Shape - Trigonous-ellipsoid
L x W - 1-1.2 cm x 0.4-0.5 cm
Color - Pink with greenish coloration
Wings - equal
Shape - Rounded to triangular
L x W - 10-12 x 5-6 mm
Number of locules - 3
Type of placentation - Axile
Style - 3
Length - 4-5 mm
Stigma - Bifid, apically forked, spiral band and
papillose all around
Color - Golden yellow
FRUIT - -
Shape Rounded Recurved horizontally
L X W 1 x 1.7 cm 10-13 mm x 5-6 mm
Color Green -
Hair Glabrous glabrous
Wings Sub-equal -
Shape Rounded -
L X W 1.2 x 0.8 cm -
Color Green -
Apex Rounded -
Base Rounded -
Hair Absent -
*green highlight = similarities, yellow highlight = updated characters, dash (“-“) = not seen or not stated in
protologue.
20
Begonia (Rubite 1001) has a rhizomatous stem and a prominent rugulose adaxial side
with a maroon colored abaxial side that contains hairs (Figure 6A & 6D). The leaves were long
ovate (Figure 6A). Majority of the Begonia were seen with pistillate and staminate flowers
(Figure 6B & 6C). This Begonia is different from the five known recorded Begonia species of
Surigao del Norte, thus it is possible that this is a new species. Therefore it was compared with
another Philippine Begonia species with rugolose leaves, Begonia obtusifolia Merr (1919).
Table 3 shows the major similarities and differences between the data collected from the
fieldwork and data from Merrill (1919).
Table 3. Major similarities and differences between Begonia sp. nov. 1 (Rubite 1000) and B. obtusifolia *
Collection Number Rubite 1001 Ramos, M. & Edano, G. 30803
Species name Begonia sp. nov. 1 Begonia obtusifolia (Merrill, 1919)
STEM Creeping creeping
Length 17 cm -
Figure 5. Begonia colorata A. Habitat; B. Fruit
21
Diameter 7.3 mm -
Internode length 11.6 mm -
Color Greenish brown -
Node Nothing in particular -
Color Greenish brown -
Hair Glabrous -
STIPULES - -
L x W 8 x 3 mm -
Shape Triangular -
Color Maroon -
Presence of keel Present -
Apex Aristate -
Hair Present -
Length 3 mm - 5 mm -
Color Light maroon -
Distribution Present in keel and in margin -
LEAVES - -
PETIOLE - Slender
Length 18.3 cm 10-13 cm
Diameter 6 mm -
Color Maroon to brown -
Hair Present -
Length 2 - 3 mm -
Color Brown -
Distribution Dense -
BLADE - -
Shape Long ovate Oblong to oblong-ovate
L x W 15.8 x 8.3 cm 7-10 x 3.5-4.5 cm
Apex Acuminate Narrowed upward to obtusely
rounded
Base Cordate Obliquely cordate
Margin Shallowly, irregularly crenate. Almost
entire. Cilliated.
Sinus shallow, narrow
Larger lobe Small lobes at the tip of each primary veins 2-4 undulations
Smaller lobe - 1 or 2 obtuse lobes
Adaxial Rugolous. Green. Some leaves are with
white islets at the interveins
Shiny
Hair Glabrous Glabrous
Abaxial Maroon colored. Trichomes at the primary
veins
Hair Present -
o Length 1 mm appressed -
o Color Maroon -
o Distribution Found only in primary veins -
INFLORESCENCE Branching 3 times Dichotomously branched at the
apex
PEDUNCLE Height of inflorescence is at level of
mature leaf or lower
-
22
Length 14.8 cm 18-24 cm
Diameter 3 mm -
Color Brownish maroon but lighter than petiole -
Hair Present Glabrous
Length 1-1.5 mm -
Color Brownish -
Distribution Less dense than the petiole -
BRACTS - -
STAMINATE FLOWER Color white -
L x W 16 x 14 mm -
PEDICEL - -
Length 7 mm -
Diameter 1 mm -
Color Pinkish green -
BRACTEOLES Present -
Shape Boat-shaped -
L x W 6 x 4 mm -
Color Light green -
TEPALS - -
OUTER - -
Number 2 -
Shape Rounded Broadly ovate
L x W 8 x 9 mm 14 x 11 mm
Color White Pink
Adaxial - -
Hair Glabrous -
Abaxial - -
Hair Glabrous -
INNER - -
Number 2 -
Shape Oblantulate Narrowly oblong
L x W 8 x 4 mm 4 mm W
Color White Pink
Adaxial Glabrous -
Hair - -
Abaxial - -
Hair Glabrous -
STAMEN - -
Union United at the base -
Filament number 35 – 40 -
Anther length 1 mm -
Shape Rounded -
Filament length 1.5 – 2mm -
Height of stamen 3 mm -
PISTILLATE FLOWER - As large as staminate flowers
L x W 16 x 10 mm -
23
PEDICEL - -
Length 11 mm -
Diameter 1 mm -
Color Greenish brown Pink
Hair Glabrous -
BRACTEOLES - -
Shape Ovate -
L x W 5 x 2 mm -
Color Green -
TEPALS - -
OUTER - -
Number 2 -
Shape Obovate to truncate -
L x W 8 x 10 mm -
Color Pinkish white Pink
Adaxial - -
Abaxial - -
INNER - -
OVARY - -
Shape Oblong -
L x W 9 x 4 mm 14 x 10 mm
Color Green -
Wings Unequal Equal
Shape Triangular -
L x W 8-9 x 2-3 mm -
Color Green -
Apex Truncate Truncate
Base Rounded Rounded
Hair Glabrous -
Number of locules 3 -
Type of placentation Axillary -
Stigma Twisted -
Color Yellow green -
FRUIT - -
Shape Obliquely oblong -
L X W 11 x 12.3 cm -
Color Brown -
Hair Absent -
Wings Un-equal -
Shape Some are rounded some are triangular -
L X W 12 x 4 mm -
Color Brown -
Apex Pointed -
Base Rounded -
Hair Absent -
*green highlight = similarities, yellow highlight = updated characters, dash (“-“) = not seen or not stated in
protologue.
24
Figure 6. Begonia sp. nov. 1 A. Habitat; B. Pistillate flower; C. Staminate flower; D. Adaxial and Abaxial side; E.
Fruit
25
Begonia (Rubite 1002) was numerous upon discovery (Figure 7A). This Begonia is
different also from the five known Begonia species of Surigao del Norte, thus it is possible that
this is also a new species. Its distinguishing character is its peltate-shaped leaf thus it was
initially compared to Begonia elmeri Merr (1918) (Figure 7B). It has a green adaxial side while
its abaxial side is maroon (Figure 7A & 7B). The color of the petiole ranges from green at its
apex up to almost brown on its base (Figure 7B). All the reproductive parts were seen upon
collection (Figure 7C, 7D & 7E). Table 4 shows the major similarities and differences between
the data collected from the fieldwork and data from Merrill (1918).
Table 4. Major similarities and differences between Begonia sp. nov. 2 (Rubite 1002) and B. elmeri *
Collection Number Rubite 1002 Elmer, A.D.E. 14183
Species name Begonia sp. nov. 2 Begonia elmeri (Merrill, 1918)
STEM Rhizomatous Succulent, curved
Length 13 cm “short”
Diameter 1.2 cm 1.25 cm
Internode length 5 - 10 mm -
Color Some green, some brown Deep red
Node - -
Color Brown -
Hair Glabrous -
STIPULES - -
L x W 19 x 13 mm 1-1.5 cm L
Shape Triangular Broadly oblong
Color Red -
Presence of keel Present. Keel possess 1.5 mm wing like
structure
-
Apex Acuminate Mucronate
Hair Present Glabrous
Length 0.4 cm -
Color Red -
Distribution Near the base of the keel -
LEAVES - Alternating
PETIOLE Height of inflorescence towers mature leaf Ascending
Length 25.5 cm 10-20 cm
Diameter 8 mm
Color Green becoming brownish to the base Deep red
Hair Glabrous Dense
BLADE - Descending, leathery
Shape Wide ovate Ovately rotund to rotund
26
L x W 11.5 x 13 cm 8-23 x 5.5-17 cm
Apex Acuminate Short, sharp point
Base Peltate Peltate
Margin Cilliated. Shallowly undulate. Shallow
toothed at the tip of the primary veins
Entire
Adaxial - Dark green, shining
Hair Glabrous Glabrous
Abaxial - Reddish brown
Hair Glabrous Glabrous
INFLORESCENCE Branching 3 times -
PEDUNCLE Swelling at the base of the peduncle Ascending from uppermost leaf
axils, subterete
Length 20 cm -
Diameter 3.5 mm “Very slender”
Color Green becoming reddish brown at the base Brown
Hair Glabrous Glabrous
BRACTS Absent Present
STAMINATE FLOWER - -
L x W 16 x 8 mm
PEDICEL - -
Length 5 mm 1 cm
Diameter 1 mm -
Color White Brown
Hair Glabrous -
BRACTEOLES Present Present
Shape Ovate -
L x W 4 x 1 mm -
Color Green -
TEPALS - -
OUTER - -
Number 2 -
Shape Obovate to truncate Orbicular
L x W 12 x 8 mm 1x1 cm
Color White Light pink
Adaxial - -
Hair Glabrous -
Abaxial - -
Hair Glabrous -
INNER - -
Number 2 -
Shape Obovate to truncate Orbicular
L x W 8 x 4 mm 1 x 1 cm
Color White Light pink
Adaxial - -
Hair Glabrous -
Abaxial - -
Hair Glabrous -
27
STAMEN - -
Union United at the base -
Filament number 25-30 Numerous
Anther length 1 mm 1 mm
Shape Rounded Obcuneate or truncately short
clavate
Filament length 1 mm 1.5-2 mm
Height of stamen 1.5 mm -
PISTILLATE FLOWER - -
PEDICEL - -
Length 10 mm -
Diameter 1 mm -
Color Greenish brown -
BRACTEOLES - -
TEPALS - -
OUTER -
Number 2 -
Shape Rounded to obtuse -
L x W 10 x 8 mm -
Color Green -
INNER - -
OVARY - -
Shape Oblong -
L x W 6 x 3 mm -
Color Green -
Wings - -
Shape Rounded to obtuse -
L x W 6 x 4 mm -
Color Green -
Apex Obtuse -
Base Rounded -
Number of locules 3 -
Type of placentation Axile -
Style - -
Length 3 mm -
Stigma - -
Color Yellow-green -
FRUIT - -
L X W - 1 x 1.8 cm
Wings - One wing much larger than the other
2
L X W - 3-4mm W, larger 1 cm
Apex - Subtruncate
Base - Rounded
*green highlight = similarities, yellow highlight = updated characters, dash (“-“) = not seen or not stated in
protologue
28
Figure 7. Begonia sp. nov. 2 A. Habitat; B. Abaxial side; C. Staminate flower; D. Fruit; E. Pistillate flower
29
Begonia (Rubite 1003) were numerous and upright (Figure 8A). Its distinguishing
character is its mottled leaves thus initially identified as Begonia leucosticta (Figure 8A, 8C &
8D). The nodes were also swollen upon closer inspection (Figure 8D). All the reproductive parts
were present upon collection (Figure 8B & 8C). Table 5 shows the major similarities and
differences between the data collected from the fieldwork and data from Hughes et al. (2015-).
Table 5. Major similarities and differences between Begonia leucosticta (Rubite 1003) and B. leucosticta *
Collection Number Rubite 1003 Merrill 5274
Species name Begonia leucosticta Begonia leucosticta (Warburg, 1904)
STEM Erect erect
Length 2m -
Diameter 10 mm -
Internode length 9.7 cm -
Color Dark green -
Node Slightly swollen -
Color Becoming brown -
Hair Glabrous Glabrous
STIPULES - -
L x W 2.5 x 0.5 cm -
Shape Lanceolate -
Color Green -
Presence of keel Present but not pronounce -
Apex Acute -
Hair Glabrous -
LEAVES - -
PETIOLE - -
Length 4.9 cm 1-4 cm
Diameter 2.5 mm -
Color Green to dark green -
Hair Glabrous Few
Distribution - Scattered
BLADE - -
Shape Oblong Lanceolate
L x W 12.7x 8 cm 12-17 x 4-5 cm
Apex Acuminate Acute
Base Cordate Almost cordate, strongly oblique
Margin Dentate Undulate, slightly denticulate
Larger lobe - Almost cuneate
Smaller lobe - Rounded-auriculate
Adaxial White to silvery spots at the
interveins.
Blue-green with big scattered white spots
Hair Glabrous Glabrous
Abaxial - -
Hair Glabrous Glabrous
INFLORESCENCE
A pair of female flowers below a
rachis of male flowers. Protogynous.
Inflorescence is opposite of a leaf
-
30
PEDUNCLE - -
Length 3.5 cm 1 cm
Diameter 2.3 mm 1 mm
Color Light green -
Hair Glabrous -
BRACTS Absent -
STAMINATE
FLOWER
Branching 4 times Compound cymose
L x W 12 x 5 mm -
PEDICEL - -
Length 7 mm 1-2 mm
Diameter 0.9 mm -
Color White -
Hair Glabrous -
BRACTEOLES - -
Shape Lanceolate -
L x W 8 x 2 mm -
Color Light green -
TEPALS - -
OUTER - -
Number 2 2
Shape Rounded orbicular
L x W 4 mm x 4 mm -
Color Pink -
Adaxial Nothing in particular -
Hair Glabrous -
Abaxial Nothing in particular -
Hair Glabrous -
INNER - -
Number 2 -
Shape Rounded -
L x W 4 mm x 4 mm -
Color Pink -
Adaxial Glabrous -
Abaxial Glabrous -
STAMEN - -
Union United at the base -
Filament number 20 -
Anther length 1 mm -
Shape Rounded -
Filament length 1.5 mm -
Height of stamen 4 mm -
PISTILLATE
FLOWER
- -
L x W 2.5 cm x 1.2 cm -
PEDICEL - -
Length 5 cm -
Diameter 1 mm -
Color Light green -
Hair Glabrous -
BRACTEOLES - -
Shape Lanceolate -
L x W 10.5 x 3 mm -
31
Color Light green -
TEPALS - -
OUTER - -
Number 2 -
Shape Ovate -
L x W 12 mm x 11 mm -
Color Pink -
Adaxial Nothing in particular -
Hair Glabrous -
Abaxial Nothing in particular -
Hair Glabrous -
INNER - -
Number 3 -
Shape Ovate -
L x W 10.5 x 5.5 mm -
Color Pink -
Adaxial Nothing in particular -
Hair Glabrous -
Abaxial Nothing in particular -
Hair Glabrous -
OVARY - -
Shape Oblong -
L x W 1.2 x 1.1 cm -
Color Pinkish green -
Wings Sub equal. Surrounding and extending
beyond ovary
-
Shape Rounded at the margins -
L x W 12 mm x 14 mm -
Color Pink -
Apex Truncate -
Base Rounded -
Hair Glabrous -
Number of locules 3 -
Type of placentation Axillary -
Style Present -
Length 4 mm -
Stigma 3 twisted -
Color Yellow -
FRUIT - -
Shape Obtriangular -
L X W 6 cm x 3.4 cm -
Color Brown -
Hair Glabrous -
Wings Present -
Shape Rounded -
L X W 2.6 x 1.6 cm -
Color Brown -
Apex Truncate -
Base Rounded -
Hair Absent -
*green highlight = similarities, yellow highlight = updated characters, dash (“-“) = not seen or not stated in
protologue
32
Figure 8. Begonia leucosticta A. Habitat; B. Abaxial side & Pistillate Flower; C. Staminate Flower; D. Stipule
33
Chapter V
Discussion
Begonia (Rubite 999) was the first species of Begonia found in Surigao del Norte. It was
confirmed B. acuminatissima through the use of the protologue by Merrill (1911). B.
acuminatisima can be distinguished from other Begonia species as a creeping Begonia having
narrowly ovate and prominently acuminate leaves. Begonia (Rubite 999) had the same kind of
long acuminate leaf tip (about 1 cm) as B. acuminatissima, prompting the researchers to identify
it as B. acuminatissima. Another similarity between the two is the presence of wings that are sub-
equal. Other data that support its identification include the same anther length, outer tepal
number, color of the staminate flower, and similarity of flowers. As seen in Table 1 B.
acuminatissima has a rhizomatous stem and 2+2 tepals (2 outer and 2 inner), species typical of
Begonia section Baryandra (Rubite et al., 2013), further strengthening the evidences that support
its identity.
B. acuminatissima was seen in different areas of Barangay Motorpool, Municipality of
Tubod. It was found at the rock crevices of a karst formation. Going higher up the trail, the same
species was also found. It was also found along roadside thriving in limestone formations. This
reflects its natural characteristics as a widespread species (Rubite, 2013). The type locality of B.
acuminatissima is Balut Island. Other areas where B. acuminatissima was recorded include
Camiguin Island, Agusan del Norte (Mt. Urdaneta and Cabadbaran.). The discovery of B.
acuminatissima in Surigao del Norte therefore provides a new record of this species in the
province. As a widespread species, the populations that have been encountered during the
fieldwork had complete reproductive and vegetative parts, enabling the researchers to completely
describe them.
34
Two records of Begonia in Surigao del Norte
Two Begonia species identified by Merrill (1911) and by Warburg (1904) were found in
Surigao del Norte, namely, Begonia bolsteri and Begonia leucosticta. B. bolsteri was the second
Begonia found during the expedition in Barangay Motorpool. Prior to the fieldwork however was
the publication by Naive et al. (2018) declaring B. bolsteri a synonym of B. colorata. Both
Begonia (Rubite 1000) and B. colorata collected by Naive et al. (2018) had the same shape, apex
and base of the leaves. Begonia (Rubite 1000) also had the same distinct dull green lamina with
distinct pale grey green patterns at the midrib of B. colorata. Despite the lack of reproductive
parts the presence of the fruits with the same equal wings which are rounded to triangular helped
greatly in the comparison of the two. Although a further study of the reproductive parts of
Begonia (Rubite 1000) is needed, in this study the researchers would like to claim that they had
found a new record of Begonia colorata.
In the Municipality of Socorro B. leucosticta was found. Since the protologue of B.
leucosticta from Perkins (1904) was not available, comparison with the herbarium specimen
(Hughes et al., 2015-) was made. Begonia (Rubite 1003) had the same small staminate flowers
and mottled large glabrous leaves of Begonia leucosticta. Dagami, Isabela is the type locality of
B. leucosticta. thus the current study therefore is reporting this new record for Surigao del Norte
Two possible new Begonia species under section Baryandra
In addition to the three previously mentioned Begonia species, there are two Begonia
species that are suspected to be new species. These two Begonia species may fall under section
Baryandra, mainly due to their rhizomatous habit, which is common for species under this
section (Rubite et al., 2013)
35
Begonia (Rubite 1001), designated as Begonia sp. nov. 1, was distinct by its rugulose
adaxial leaf surface. Since there is no record of rugolose Begonia species in Mindanao Begonia
(Rubite 1001) was compared with another Philippine rugolose B. obtusifolia (Merrill, 1919),
Both are are characterized by rhizomatous stem, persistent stipules, four tepals and inferior
ovary. All of these indicate that the two belong to Begonia section Baryandra. But they also
differ significantly, Begonia sp nov 1 leaf shape is long ovate with long acuminate apex while B.
obtusifolia is oblong to oblong-ovate in shape and with obtusely rounded apex. Another
difference is that hairs are present in the peduncle of B. sp. nov. 1 and the color of the pedicel of
its pistillate flower is entirely different when compared with the descriptions in the protologue of
B. obtusifolia. Lastly, the ovary of B. sp nov 1 is with unequal wings while that of the B.
obtusifolia are equal. Thus, Begonia (Rubite 1001) is proposed as new species.
Begonia (Rubite 1002), designated as Begonia sp. nov. 2, was distinct by its peltate leaves.
Thus it was compared to another peltate species from Mindanao, Begonia elmeri (Merrill, 1918).
B. sp. nov. 2 was cross-examined with the protologue of B. elmeri. Similarities between the two,
include the shape of the leaf blade; the almost entire margin, and the glabrous blade and
peduncle. However, differences between the two were also observed such as B. sp. nov. 2 having
shallowly undulated margin with hairs present along it as compared to B. elmeri having an entire
margin. B. sp. nov. 2 has a white colored peduncle as opposed to the brown colored peduncle of
B. elmeri. There is a possibility that the two species are the same. However, the differences
between the two, the incompleteness of important details in the protologue of Merrill (1918)
such as descriptions about the staminate and pistillate flowers, and the absence of fruits of B sp.
nov. 2, limits the cross-examination between the two species and validating their similarity. This
36
prompts the researchers to designate B. sp nov 2 to be a potential novel species until a complete
morphological data is collected, or molecular data would prove otherwise.
The potential new species require confirmations since morphological characters do not
always validate novel discoveries. Additionally, not all of the possible new species have
complete morphological data. Without a complete morphological data, the evidence supporting
the novelty of these Begonia species is weak. The best way to prove that they are new species
and not just variants is by using molecular data such as chloroplast DNA (Peng et al., 2017a;
Pereira et al., 2008) in addition to other evidences. This does not disregard the usefulness of
morphological data as they serve as baseline information for researchers.
37
Chapter VI
Conclusion and Recommendation
The province of Surigao del Norte remains understudied. Due to this fact, there has been
no update about the Begonia species of the province. However, through the botanical
exploration, a taxonomic update of the Begonia species of Surigao del Norte was produced. A
total of five Begonia species had been identified, namely, B. acuminatissima, B. colorata, B.
leucosticta and two possible novel species. The recovery and discovery of these five Begonia
species reflects the rich biodiversity of Surigao del Norte, and the Philippines as a whole. It also
shows that the primary forests of the Philippines are still not fully explored botanically, as many
species are just waiting to be discovered.
This continuous discovery of different Begonia species in the Philippines further
reinforces our status as one of the centers of mega-biodiversity. Though different species are
recovered throughout the years, it cannot be denied that they are always accompanied with
threats that may continue without proper conservation of primary forests. Promoting awareness
for the Begonia would help secure a more stable future for the propagation of these highly
endangered species.
Despite the recovery of the said species of Begonia, more studies are still needed to
overcome the lack of research over the past decades. It is recommended that the areas explored
be revisited in order to complete the lacking morphological characters for the different species. A
complete set of morphological characters would make it possible to update phylogenetic
relationships within the different species of Begonia within Surigao del Norte.
38
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http://surigaonorte.tripod.com/introphysical.htm#soil
Tacio, H. (2009). Philippines’ Ecosystem on the Verge of Extinction. Retrieved from:
https://www.gaiadiscovery.com/nature-biodiversity/philippines-ecosystems-on-the-verge-
of-extinction.html/
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Appendix A
Wildlife Gratuitous Permit of Mr. Mark Gregory Q. Rule
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