LIANA CLIMBING STRATEGIES AND RELATIONSHIP WITH HOST …

LIANA CLIMBING STRATEGIES AND RELATIONSHIP WITH HOST TREES IN MO SINGTO FOREST DYNAMICS PLOT,

KHAO YAI NATIONAL PARK, THAILAND

NATTHIDA KHIEWBANYANG

A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR

THE DEGREE OF MASTER OF SCIENCE (ENVIRONMENTAL BIOLOGY)

FACULTY OF GRADUATE STUDIES MAHIDOL UNIVERSITY

2015

COPYRIGHT OF MAHIDOL UNIVERSITY

Thesis entitled



……………….………….…..………….. Miss Natthida Khiewbanyang Candidate

……………………….….…..………… Asst. Prof. Jenjit Khudamrongsawat, Ph.D. (Biology) Major advisor

……………………….….…..………….. Lect. Chanpen Saralamba, Ph.D. (Biology) Co-advisor

……………………….….…..………. Prof. Patcharee Lertrit, M.D., Ph.D. (Biochemistry) Dean Faculty of Graduate Studies Mahidol University

……………….….…..………………….. Assoc. Prof. Prayad Pokethitiyook, Ph.D. (Chemical Engineering) Program Director Master of Science Program in Environmental Biology Faculty of Science Mahidol University

Thesis entitled



was submitted to the Faculty of Graduate Studies, Mahidol University for the degree of Master of Science (Environmental Biology)

on March 9, 2015

……………….………….…..………… Miss Natthida Khiewbanyang Candidate

………………….…..…………………. Asst. Prof. Wirong Chanthorn, Ph.D. (Biology) Chair

……………………….….…..………… Prof. Patcharee Lertrit, M.D., Ph.D. (Biochemistry) Dean Faculty of Graduate Studies Mahidol University

……………………….….…..……….. Asst. Prof. Jenjit Khudamrongsawat, Ph.D. (Biology) Member ……………………….….…..………… Lect. Chanpen Saralamba, Ph.D. (Biology) Member ……………….….…..………………… Asst. Prof. Somsak Dangtip, Ph.D. (Applied Nuclear Physics) Acting Dean Faculty of Science Mahidol University

iii

ACKNOWLEDGEMENTS

I would like to express my gratitude to my advisor, Assistant Professor Dr.

Jenjit Khudamrongsawat for the valuable guidance, advice, and motivation. This

master thesis would not be possible without her continuous support and patience.

I am grateful to my committee members Dr. Chanpen Saralamba for

introducing me to the topic as well for the support, encouragements and valuable

advices through the learning process of this master thesis. I would also like to thank

Assistant Professor Dr. Wirong Chanthorn for being the external examiner of my

thesis defense including useful suggestions for improvement. My grateful thanks are

also extended to Miss. Anuttara Natalang who gave the permission to use all required

information, the necessary material to complete the research work and including useful

comments, remarks and engagements. I am using this opportunity to express my

gratitude to Professor Warren Y. Brockelman who gave me an opportunity to

participate and learn in Mo Singto Forest Dynamics Plot.

This thesis work is supported by Thailand Graduate Institute of Science

and Technology (TGIST), TGIST-01-55-020 under National Science and Technology

Development Agency (NSTDA).

I gratefully thank Department of National Parks, Wildlife and Plant

Conservation for permission to collect information in Khao Yai National Park.

Also, I would like to thank the assistants in my fieldwork, who have

willingly shared their precious time during the process of fieldwork. I thank my fellow

graduate students and friends for their various helps and supports.

Finally, I would like to thank my parents for their kindness, wonderful

support and understanding throughout my study.

Natthida Khiewbanyang

iii

iii

Fac. of Grad. Studies, Mahidol Univ. M.Sc. (Environmental Biology) / 62

BIOGRAPHY

NAME Natthida Khiewbanyang

DATE OF BIRTH 28 October 1987

PLACE OF BIRTH Bangkok, Thailand

INSTITUTIONS ATTENDED Mahidol University, 2006-2009

Bachelor of Science

(Conservation Biology)

Mahidol University, 2010-2014

Master of Science

(Environmental Biology)

SCHOLARSHIP RECEIVED Scholarship for Young Scientists,

2010-2011

Thailand Graduate Institute of Science and

Technology, 2012-2013

HOME ADDRESS 105 Soi Suan Phak 29, Suan Phak Rd.,

Taling Chan, Bangkok, THAILAND 10170

E-mail: [email protected]


APPENDIX

Natthida Khiewbanyang Appendix / 56

Table A. List of host trees species ≥ 10 cm dbh in the Mo Singto Forest Dynamics

Plot, Khao Yai National Park, Thailand

Tree speciesNumber of host trees

Number of host treeclimbed by lianas

% Liana climbing

Sloanea sigun 1008 107 8.55

Nephelium melliferum 735 96 7.67

Ilex chevalieri 1021 73 5.83

Dipterocarpus gracilis 757 56 4.47

Symplocos cochinchinensis 768 56 4.47

Mastixia pentandra 768 55 4.39

Gironniera nervosa 708 50 4.00

Lithocarpus thomsonii 364 41 3.28

Choerospondias axillaris 192 39 3.12

Cleistocalyx operculatus 414 35 2.80

Aidia densiflolia 399 27 2.16

Balakata baccata 58 25 2.00

Cinnamomum subavenium 474 25 2.00

Nauclea orientalis 227 25 2.00

Castanopsis acuminatissima 261 23 1.84

Beilschmiedia maingayi 347 22 1.76

Michelia baillonii 61 21 1.68

Aphanamixis polystachya 170 18 1.44

Pterospermum cinnamomeum 79 17 1.36

Pterospermum simiarum 269 16 1.28

Schima wallichii 164 16 1.28

Syzygium syzygioides 152 16 1.28

Cinnamomum ilicioides 90 15 1.20



Plot, Khao Yai National Park, Thailand (Cont.)



% Liana climbing

Gonocaryum lobbianum 590 15 1.20

Alphonsea boniana 56 14 1.12

Beilschmiedia glauca 327 14 1.12

Dysoxylum cyrtobotryum 203 14 1.12

Knema elegans 479 14 1.12

Aquilaria crassna 239 13 1.04

Gracinia benthamii 113 12 0.96

Chionanthus microstigma 14 12 0.96

Helicia formosana 168 11 0.88

Prunus javanica 105 11 0.88

Sarcosperma arboreum 191 11 0.88

Bridelia insulana 78 10 0.80

Walsura robusta 258 10 0.80

Mischocarpus pentapetalus 72 8 0.64

Platea latifolia 152 8 0.64

Carallia brachiata 34 7 0.56

Phoebe lanceolata 96 7 0.56

Adinandra integerrima 59 6 0.48

Ficus nervosa 29 6 0.48

Schefflera heptaphylla 104 6 0.48

Syzygium grande 53 6 0.48

Eurya nitida 93 5 0.40

Excoecaria oppositifolia 91 5 0.40






% Liana climbing

Fraxinus floribunda 33 5 0.40

Lithocarpus aggregatus 58 5 0.40

Machilus odoratissima 97 5 0.40

Podocarpus neriifolius 37 5 0.40

Polyosma elongata 218 5 0.40

Aglaia elaeagnoidea 211 4 0.32

Aglaia odoratissima 104 4 0.32

Palaquium garrettii 48 4 0.32

Syzygium siamensis 98 4 0.32

Canarium subulatum 35 4 0.32

Chukrasia tabularis 29 4 0.32

Erythrina subumbrans 18 3 0.24

Litsea beusekomii 104 3 0.24

Litsea verticillata 106 3 0.24

Morus macroura 8 3 0.24

Neolitsea zeylanica 25 3 0.24

Rhus rhetsoides 46 3 0.24

Sandoricum koetjape 19 3 0.24

Alchornea rugosa 80 3 0.24

Altingia excelsa 11 3 0.24

Anthocephalus chinensis 14 3 0.24

Apodytes dimidiata 45 3 0.24

Ardisia sanguinolenta 54 2 0.16






% Liana climbing

Baccaurea ramiflora 28 2 0.16

Diospyros glandulosa 27 2 0.16

Gomphandra tetrandra 87 2 0.16

Lindera communis 38 2 0.16

Litsea monopetala 7 2 0.16

Litsea salicifolia 26 2 0.16

Memecylon edule 75 2 0.16

Miliusa lineata 32 2 0.16

Toona ciliata 24 2 0.16

Turpinia nepalensis 16 2 0.16

Aglaia edulis 11 2 0.16

Aglaia lawii 34 2 0.16

Antiaris toxicaria 19 2 0.16

Aphananthe cuspidata 20 2 0.16

Archidendron lucidum 13 2 0.16

Ardisia nervosa 37 2 0.16

Beilschmiedia affintermedia 55 2 0.16

Buchanania arborescens 4 1 0.08

Chionanthus ramiflorus 39 1 0.08

Chisocheton grandiflorus 11 1 0.08

Claoxylon indicum 7 1 0.08

Dendrocnide stimulans 8 1 0.08

Elaeocarpus petiolatus 6 1 0.08






% Liana climbing

Eugenia grata 16 1 0.08

Ficus glaberrima 10 1 0.08

Ficus stricta 5 1 0.08

Ficus vasculosa 17 1 0.08

Hopea odorata 1 1 0.08

Hydnocarpus ilicifolius 2 1 0.08

Ilex affchapaensis 3 1 0.08

Lithocarpus pattaniensis 25 1 0.08

Macaranga denticulata 1 1 0.08

Macaranga siamensis 18 1 0.08

Machilus affsalicina 8 1 0.08

Mallotus paniculatus 1 1 0.08

Mangifera longipetiolata 25 1 0.08

Markhamia stipulata 4 1 0.08

Nothapodytes montana 37 1 0.08

Ormosia sumatrana 13 1 0.08

Phoebe cathia 12 1 0.08

Platymitra macrocarpa 22 1 0.08

Podocarpus imbricatus 3 1 0.08

Pouteria stellibacca 3 1 0.08

Prunus arborea 28 1 0.08

Saurauia roxburghii 57 1 0.08

Shorea henryana 2 1 0.08






% Liana climbing

Sterculia balanghas 5 1 0.08

Tetrameles nudiflora 18 1 0.08

Triadica cochinchinensis 4 1 0.08

Viburnum cylindricum 50 1 0.08

Xanthophyllum virens 18 1 0.08

Total 15620 1251

Fac. of Grad. Studies, MahidolUniv. Thesis / iv

LIANA CLIMBING STRATEGIES AND RELATIONSHIP WITH HOST TREES IN MO

SINGTO FOREST DYNAMICS PLOT, KHAO YAI NATIONAL PARK, THAILAND

NATTHIDA KHIEWBANYANG 5337755SCEB/M

M.Sc. (ENVIRONMENTAL BIOLOGY)

THESIS ADVISORY COMMITTEE: WIRONG CHANTHORN, Ph.D.

JENJIT KHUDAMRONGSAWAT, Ph.D., CHANPEN SARALAMBA, Ph.D.

ABSTRACT

Lianas are woody climbing plants exhibiting a unique habit that requires host

trees as physical supports. High diversity of lianas in tropical forests involves its various and

distinct climbing strategies to ascend to the forest canopies in addition to species richness. The

diverse tactics of its climbing strategies are dependent on variations in the host tree

characteristics, particularly, the sizes and bark textures of host trees. The perennial plant

characteristics of trees on 30.5 hectares of the Mo Singto Forest Dynamics Plot, which is a

seasonal evergreen forest located in the Khao Yai National Park in Thailand, were examined.

Thirty ramets per liana species with diameter at breast height ≥ 3 cm of 1,560 liana ramets

were examined, and six climbing strategies were classified in this study. The dominant

climbing strategy was twiners, followed by the combination of twiners and scramblers,

scramblers, tendril, hook and adventitious root climbers, respectively. Liana ramet size was

positively correlated with the host tree size. However, this relationship varied with the type of

climbing strategies. Most twiners, hooks and tendril climbers tended to use small host trees.

The scrambler and the combination of twiner and scrambler ramets appeared to be

independent of host tree sizes. Adventitious root climbers tended to use large host trees. Most

ramets used host trees with slightly rough bark texture more often than other types of bark

texture with the exception of the adventitious root climbers that were often found on trees with

rough bark texture. In conclusion, the presence and abundance of ramets with different

climbing strategies could be associated with sizes and bark textures of host trees.

KEY WORDS: LIANAS / CLIMBING STRATEGY / SEASONAL EVERGREEN

FOREST / KHAO YAI NATIONAL PARK / HOST TREES

62 pages

Fac. of Grad. Studies, MahidolUniv. Thesis / v

กลวิธีการเลี้อยของเถาวัลยและความสัมพันธกับพืชใหอาศัยในแปลงสํารวจถาวรมอสิงโต อุทยานแหงชาติเขาใหญLIANA CLIMBING STRATEGIES AND RELATIONSHIP WITH HOST TREES IN MO SINGTO FOREST DYNAMICS PLOT, KHAO YAI NATIONAL PARK, THAILAND

ณัฐธิดา เขียวบานยาง 5337755 SCEB/M

วท.ม. (ชีววิทยาสภาวะแวดลอม)

คณะกรรมการที่ปรึกษาวิทยานิพนธ: วิรงค จันทร, Ph.D., เจนจิต คูดํารงสวัสดิ์, Ph.D., จันทรเพ็ญ ศรลัมพ, Ph.D.

บทคัดยอเถาวัลยหรือไมเถาเนื้อแข็งเปนพืชที่มีวิสัยเฉพาะตัวโดยมีโครงสรางของลําตนทอดเลื้อยหรือปน

ปายตนไมอื่นเพื่อยึดลําตนใหทรงตัวอยูได ความหลากชนิดของเถาวัลยในปาเขตรอนยังเกี่ยวของกับความหลากหลายของกลวิธีการเลื้อยที่เถาวัลยใชสําหรับเคลื่อนที่ไปสูเรือนยอด ความหลากหลายของกลวิธีการเลื้อยดังกลาวอาจขึ้นอยูกับลักษณะของพืชใหอาศัยโดยเฉพาะขนาดและลักษณะเปลือก จึงทําการศึกษาความสัมพันธดังกลาวบริเวณปาดิบชื้นแบบมีฤดูกาลในแปลงสํารวจถาวรขนาดพื้นที่ 30.5 เฮกตาร โดยเลือกศึกษาจากเถาวัลย 52 ชนิด ชนิดละ 30 ตน ขนาดเสนผาศูนยกลางลําตนมากกวาหรือเทากับ 3 เซนติเมตร จํานวนทั้งสิ้น 1,560 ตนและจําแนกกลวิธีการเลื้อย จากการศึกษาสามารถจําแนกกลวิธีการเลื้อยไดหกวิธี คือ การใชลําตนหรือกิ่งเลื้อยพัน (Twiners) การใชลําตนหรือกิ่งเลื้อยพันและพาดพิง (The combination of twiners and scramblers) การเลื้อยพันแบบพาดพิง (Scramblers) การใชมือพัน (Tendrils) การใชขอเกี่ยว (Hooks) และการใชรากพิเศษ (Adventitious root climbers) นอกจากนี้ยังพบวาขนาดของเถาวัลยมีความสัมพันธกับขนาดของตนไม แตอยางไรก็ดีความสัมพันธนี้ยังขึ้นอยูกับกลวิธีการเลื้อยที่เถาวัลยเลือกใชดวย โดยเถาวัลยที่ใชลําตนหรือกิ่งเลื้อยพัน ใชขอเกี่ยวและใชมือพันมักจะปรากฏบนตนไมที่มีขนาดเล็ก เถาวัลยที่ใชลําตนหรือกิ่งเลื้อยพันและพาดพิงและการเลื้อยพันแบบพาดพิงมักจะไมขึ้นอยูกับขนาดของตนไม และเถาวัลยที่ใชรากพิเศษมักจะปรากฎบนตนไมที่มีขนาดใหญ เถาวัลยสวนใหญจะปรากฎบนตนไมที่มีเปลือกคอนขางขรุขระยกเวนเถาวัลยที่ใชรากพิเศษมักจะปรากฎอยูบนตนไมที่มีเปลือกขรุขระมาก ในการศึกษาครั้งนี้แสดงใหเห็นวาการปรากฎและจํานวนของเถาวัลยที่ใชกลวิธีการเลื้อยตางๆกันมีความสัมพันธกับขนาดและลักษณะของเปลือกของพืชใหอาศัย

62 หนา

vi

CONTENTS

Page

ACKNOWLEDGEMENTS iii

ABSTRACT (ENGLISH) iv

ABSTRACT (THAI) v

LIST OF TABLES viii

LIST OF FIGURES ix

CHAPTER I INTRODUCTION 1

CHAPTER II LITERATURE REVIEWS 4

2.1 Lianas 4

2.2 Liana abundance and diversity 5

2.3 The roles and functions of lianas 6

2.4 Liana climbing strategies 6

2.5 Liana climbing stratrgies and their limiting factors 7

CHAPTER III MATERIALS AND METHODS 10

3.1 The Mo Singto Forest Dynamics Plot 10

3.2 Liana database and sampling 13

3.3 Liana climbing strategies, liana and host tree sizes, 14

and host tree bark textures

3.3.1 Climbing strategy classification 14

3.3.2 Liana and host tree size classification 19

3.3.3 Bark texture classification 20

3.4 Data anlyses 24

3.4.1 Liana climbing strategies 24

3.4.2 Relationships between liana size and host tree size, 24

liana climbing strategies and host tree size,

and liana climbing strategies and bark texture

vii

CONTENTS (cont.)

Page

CHAPTER IV RESULTS 26

4.1 Liana and host tree abundance 26

4.2 Liana climbing strategies 29

4.3 Liana and host tree relationship 31

4.3.1 Relationship between liana ramet sizes 31

and host tree sizes

4.3.2 Relationship between liana climbing strategies 33

and host tree sizes


and different host tree bark textures

CHAPTER V DISCUSSION 41

5.1 Lianas and host tree abundance 41

5.2 Lianas climbing strategies 41

5.3 Liana and host tree relationships 44

5.3.1 Relationship between liana ramet sizes 44

and host tree sizes


and host tree sizes


and host tree bark textures

REFERENCES 49

APPENDIX 55

BIOGRAPHY 62

Natthida Khiewbanyang Materials and Methods / 12

Figure 3.2 Map of the terrain on the Mo Singto Forest Dynamics Plot showing the

sampling area of 30.5 ha in four different habitats (hilltop, valley, slope and lowland)

and 20-meter survey points, the elevation ranging from 725-815 m above sea level

with 5-meter contour intervals, and the scale displaying distances on the axes in meters

(Brockelman et al., 2011).

m


3.2 Liana database and sampling

The data in this study obtained from liana and tree censuses managed by

the Ecology Laboratory of the National Center for Genetic Engineering and

Biotechnology (BIOTEC), under the National Science and Technology Development

Agency (NSTDA). Inventories of lianas and trees were kept in the form of computer

database using Microsoft Access with the following information: all living lianas and

tree stem diameter at breast height (dbh), species identification, and their locations that

were plotted into the x and y coordinate system. For this study, the 2011 recensus data

were used for liana ramet sampling.

A liana genet was defined as a rooted stems arising from a vegetative stem

of a plant. It could climb on a host trees but sometimes fell back to the ground. A

ramet (distinct genetic individual) ≥ 3 cm dbh was defined as an independent

individual that was not connected to any other aboveground or belowground stems

(Gerwing, 2004).

Trees were considered to be hosts when liana ramets climbed the trunk or

branch of the trees and were classified as primary host trees. They were the initial

support (Nabe-Nielsen, 2001).

All liana ramets with their host trees were assigned specific tagging

numbers. Monocotyledons, specifically rattans, that were the only monocot species

found in the study site and some hemiepiphytes were excluded because they were

represented by very few species and low in abundance. Lianas that could not

confidently be identified were also excluded from analyses. All selected liana ramets

were verified their living status to confirm the record in the database.

This study focused on the common species of liana ramets. All living

species were divided into common and rare species based on their densities (number

of ramets per hectare). Common species were species found at least one individual per

hectare while rare species were those with less than one individual per hectare. Thirty

ramets for each common species of lianas whose diameter at breast height (dbh) were

greater than or equal to 3 cm and found to climb host trees were randomly selected

from their tagging numbers by using a computer program (RAND function in

Microsoft office 2007). Thus, a total of 1,560 ramets of common liana species were

examined.


3.3 Liana climbing strategies, liana and host tree sizes, and host tree

bark textures

3.3.1 Climbing strategy classification

The climbing strategies for each liana stem were classified to five major

categories, which were tendril climbers, root climbers, twiners, scramblers, and hook

climbers as described in Chittibabu & Parthasarathy (2001); Dewalt et al. (2000),

Isnard & Silk (2009); Padaki et al. (2000); Putz (1984); Senbeta et al. (2005) (Figure

3.3). Other climbing strategies that were not included in these categories were also

described and recorded if found in the plot. All lianas encountered were photographed

for species confirmation.

Tendril climbers refer to modified leaves, leaflets or stipules, which twine

around a support (Chittibabu & Parthasarathy, 2001; Putz, 1984). Tendril grasp only

small diameter supports.

Root climbers refer to adventitious roots or aerial roots, adhered to host

trees and growing into cracks and crevices of the bark (Putz, 1984; Padaki et al.,

2000).

Twiners refer to apical portion of the shoots, branches or petioles, which

twine around the host trees (Dewalt et al., 2000; Isnard & Silk, 2009).

Scramblers refer to sprawlers, which lean on but do not closely attach to

host trees (Chittibabu & Parthasarathy, 2001; Putz, 1984).

Hook climbers refer to an organ such as recurved spines, hooks, or thorns,

which sprawl and attach to host trees (Isnard & Silk, 2009; Pakadi et al., 2000).


Figure 3.3 The characteristics of a tendril showing the stem that is modified into an

organ that grasp a host tree.

Figure 3.4 The characteristics of a hook showing the recurved spines used for

sprawling and attaching to a host tree.

Modified stem of tendril

Recurved spine ofhook climber


Figure 3.5 The characteristics of a twiner showing the shoot or branch twining around

the host tree trunk.

Shoot of twiner

Host tree trunk


Figure 3.6 The characteristics of adventitious roots showing attaching to supporting

host tree.

Adventitious roots of root climber

Host tree trunk


Figure 3.7 The characteristics of scramblers showing sprawling branches.


3.3.2 Liana and host tree size classification

Sizes of liana ramets were divided into three groups. Small size was

defined as having a dbh of 3-5.9 cm while the medium and large sizes were divided as

having a dbh of 6-8.9 cm and ≥ 9 cm, respectively.

Host trees with dbh greater than or equal to 10 cm being attached by the

selected ramets were also examined. Size of host trees were divided into ten size

classes which were 10-19.9, 20-29.9, 30-39.9, 40.49.9, 50-59.9, 60-69.9, 70.79.9, 80-

89.9, 90-99.9, and ≥ 100 (Figure 3.8). Based on their size distribution, host trees were

categorized into three groups. Small size (10-39.9 cm), medium size (40-69.9 cm) and

the large size group (≥ 70 cm).

Figure 3.8 Frequency distribution of host trees ≥10 cm dbh in the Mo Singto Forest

Dynamics Plot, Khao Yai National Park, Thailand. The three groups of host tree sizes

which were divided into small (10-39.9 cm dbh), medium (40-69.9 cm dbh) and large

(≥ 70 cm dbh).

Small size group Medium size group Large size group


3.3.3 Bark texture classification

Bark texture varied from rough to smooth, depending on characteristics

including lenticels, branch scars, and wounds. Lichens and mosses were not

considered as a part of bark texture. Bark texture of hosts (all of them are associated

with ramets) were classified into smooth (S), slightly rough (SR) and rough (R)

(Campbell & Newbery, 1993; van der Heijden et al., 2008). All host trees were

photographed for confirmation of their bark characteristics by comparing with the

photographs and references as below.

Smooth bark or unbroken bark is the surface being smooth and glossy to

dull area. Generally these have a very thin outer bark and flat (Figure 3.9 a-b).

Slightly rough bark is the surface being any combination of small

dimples, shallow fissures and lenticels visible (Figure 3.10 c-f).

Rough bark is the surface with (g) scales or plates, (h) deeply fissures or

cracks, (i) deeply furrows or ridges, and (j) vertical strips (Figure 3.4 g-j).


Figure 3.9 Variations in characteristics of the smooth bark texture of host trees (a) and

(b).

a

b


Figure 3.11 Variations in characteristics of the rough bark texture of host trees (g)-(j)

g h

ji


3.4 Data analyses

3.4.1 The climbing strategies of lianas

A type of climbing strategy (twiners, scramblers, adventitious roots,

tendrils, hooks and others if any) exhibited in each ramet was recorded.

The proportion of each climbing strategy was calculated by taking the

number of individual ramets with a given climbing strategy divided by the total

number of individual ramets of all climbing strategy.

Number of individual ramets for each climbing strategy

Total number of individual ramets

In terms of number of species, the proportion was calculated by taking the

number of ramet species with a given climbing strategy divided by the total number of

ramet species of all climbing strategy.

Number of ramet species for each climbing strategy

Total number of individual ramet species

To determine whether liana ramets demonstrate a selection for particular

host tree species, the number of individual trees observed as hosts was compared with

the number of individuals expected to be hosts using the 2 test (Chittibabu &

Parthasarathy, 2001). The average number of individual trees was calculated to

provide the expected proportion of liana colonization. The observed number of trees

hosting different number of lianas was calculated for each tree species.

3.4.2 Relationships between liana size and host tree size, liana climbing

strategies and host tree size, and liana climbing strategies and bark texture

Two different aspects of climbing strategies of lianas were examined. First,

simple linear regression analysis was conducted in order to clarify trends in the

relationship between liana ramet and host tree sizes. The dbh sizes of liana ramets and

host trees were plotted against each other. Second, the relationship between liana size

Proportion =

Proportion =


of each climbing strategy and host tree size were separately determined. The sizes

were plotted against one another with host tree dbh as the independent variable and

ramet dbh as the dependent variable.

The relationship between ramet size and host tree bark texture was

determined by plotting the number of ramets according to their size groups in a bar

graph. Each bar represented the number of ramet in an assigned size and separated into

different shades based on the types of host tree bark texture the ramet climbed. The

relationship between liana climbing strategies and host tree bark texture was examined

by plotting the number of ramets based on their size as bars. Different shades on a bar

were the number of ramets that use hosts of different bark texture. The preference of

hosts with particular bark texture was tested by using chi-square test based on the

availability of hosts.

The 2 and simple linear regression were analyzed by the PASW Statistics

for Windows, Version 18.0. Chicago: SPSS Inc. USA. All syntheses were test with

statistically significant relationships (p<0.05).

Natthida Khiewbanyang Literature Review / 4

CHAPTER II

LITERATURE REVIEW

2.1 Lianas

Climbing plants are classified into four fundamental categories (Gentry,

1991). The first category is comprised of herbaceous epiphytes, hemiepiphytes and all

herbaceous species that climb host trees by attaching their adventitious roots to host

tree trunks, e.g., Peteris pellusida and Asplenium nidus. The second category of these

climbers is woody hemiepiphytes and stranglers that begin life as epiphytic seedlings

by being rooted after reaching the ground (e.g. Ficus sagittata). Some species begin as

terrestrial climbers and use their adventitious root systems and/or losing contact for

climbing. The third category of woody climber plants is vines that are long and thin-

stemmed climbers, sometimes are referred to herbaceous vines. Many examples of

vines are found in the morning glory family Convolvulaceae. Vines emerge as

terrestrial seedling and are capable of growing in disturb habitats or forest edges. The

last category is lianas that have long and thick stems. Similar to vines, lianas emerge

as terrestrial seedling but are also able to grow in mature forests. They are also called

liane, brush rope, and woody vine (Putz, 1984). There are several common lianas in

tropical forests such as Tetracera indica and Gnetum montanum. Lianas also include

the climbing palms (the rattans).

Lianas exhibit a variety of shapes and sizes. They have evolutionary

evolved in diverse taxa such as gymnosperms, palms and monocotyledons (Cai et al.,

2009). In general, lianas have long and wide vessels that ensure efficient water

transport throughout their long stems (Cai et al., 2009; Isnard & Silk, 2009). They are

rooted in the ground but need structural support for their long and narrow stems (Putz,

1984; Putz & Chai, 1987). Lianas can reproduce by seeds or some portion of stem

such as sprouting, which enable them to colonize host trees (Nabe-Nielsen & Hall,

2002). Young lianas may not depend on host trees for structural support as the growth

rate is slow. The flexible stems of lianas allow them to grow without support up to


about 1.5 m in height. As they approach their maturity, rapid growth rate occurs, and

their stems may need host trees to ascend the canopy. Without host trees for structural

support, lianas can still continue their growth but at much slow rates (Putz 1984).

Lianas can grow long distances from the initial rooting point, and so long-distance

cronal is a general method of liana colonization (Nabe-Nielsen & Hall, 2002; Yorke et

al., 2013). Lianas are shade tolerant plants waiting suitable light conditions (Malizia &

Grau, 2006; Schnitzer & Bongers, 2002) and supports (Campanello et al., 2007; Carse

et al., 2000; Malizia & Grau, 2006; Ladwig & Meiners, 2010). Upon reaching host

trees, lianas increase their growth rates, sizes and reproductive outputs. Lianas are able

to survive when they fall from their host trees (Peñalosa, 1984 but see Putz, 1984).

Lianas can produce physiologically independent ramets making liana genets very long

lived. A capacity of liana genets to survive by reproducing asexually when they are

unable to regenerate sexually may also enable them to survive in a wider range of

habitats than do other plants.

2.2 Liana abundance and diversity

Lianas diversity throughout the world had differences in terms of overall

abundance and richness. In the temperate and subtropical forests, lianas are common

in many areas but their abundance and richness are lower than those in tropical forests

(Durigon et al., 2014; Schnitzer & Bongers, 2002). In tropical forests, liana

abundance, diversity, and taxonomic composition are relatively high among tropical

regions especially in Africa (Senbeta et al., 2005). Many similar studies in different

tropical forests have reported similar results (e.g. carried out in studies Addo-Fordjour,

2008; Muoghalu & Okeesan, 2005; Muthuramkumar & Parthasarathy, 2000; Putz,

1984; Putz & Chai, 1987; Reddy & Parthasarathy, 2003; Schnitzer et al., 2012).

Although lianas are abundant common in many temperate forests, abundance,

diversity and taxonomic composition is low compared with tropical forests (Schnitzer

et al., 2000). A review of liana diversity inventories in various tropical forest sites is

presented in Table 2.1.


2.3 The roles and functions of lianas

Lianas not only form a community component but also play important

roles in several aspects of forest dynamics such as suppressing tree regeneration

(Schnitzer & Carson, 2010), increasing tree mortality (Ingwell et al., 2010), serving as

food sources for many animals such as hornbills, monkeys and gibbons (Kitamura et

al., 2004; Kunz & Linsenmair, 2010; Vogel et al., 2009), providing habitat corridors

for arboreal animals such as gibbons and monkeys Schnitzers & Bongers, 2002) and

binding tree causing the stability of the individual trees (Schnitzers & Bongers, 2002).

The influence of lianas on forest dynamics may result from their interactions with host

trees. Despite many advantages of lianas, forest structures may be modified if liana

diversity and abundance change (DeWalt et al., 2000). In areas where lianas become

abundant, they may negatively impact tree species in gaps and disturbance areas

(Mohandass et al., 2015). For instance, lianas can obstruct tree growth and

regeneration by competing for space (Schnitzer et al., 2000) and causing mechanical

damages such as removing bark, shoot and bud of host trees (Putz, 1984). However,

this negative effect benefits pioneer tree species by promoting growth, recruitment and

survival of these trees (Schnitzer & Carson, 2010).

2.4 Liana climbing strategies

Most lianas climb their hosts using different climbing strategies and

require suitable structural supports to move upward. Climbing plants also show great

diversity in their climbing strategies. Liana climbing strategies have been classified

into tendril climbers, root climbers, twiners, scramblers and hook climbers

(Anbarashan & Parthasarathy, 2013; Campanello et al., 2007; Chittibabu &

Parthasarathy, 2001; DeWalt et al., 2000; Putz, 1984; Senbeta et al., 2005).

Twiners are portions of shoots, branches, or petioles that twine or coil

around supports. Twiners often climb in one direction, and show a predisposition to

turn to the right (Burnham & Revilla-Minaya, 2011). The twiner strategy is efficient

for ascending small host trees. This mode of attachment requires close contact with

surface. The main stem twines around support and is replaced by lateral branches.


Lianas develop an extensive root system. Root climbers use an

adventitious root to attach to the bark of host trees. This mode of attachment requires

close contact with surface (Putz, 1984). Tendril climbers are the modified leaves,

leaflets, inflorescence, flower peduncle or stipules twine around a support (Chittibabu

& Parthasarathy, 2001, Putz, 1984). At their young state, tendrils are relatively straight

and sensible to touch. Rapid growth occurs when tendrils come into contact with host

trees. Tendril climbers can grasp small supports. They are more suitable to gaps and

forest edges where small supports are usually more common than in forest interiors.

Hook climbers are the passive climbing strategy (Pakadi et al., 2000). This

strategy uses an organ such as recurved spines, hooks, thorns, or outgrowth as a

prickle to attach to the surface of host trees (Putz, 1984). Scramblers or sprawlers lean

on but do not become closely attach to host trees (Putz, 1984). Sometimes, the

scramblers use leader shoots to climb to the host trellis and other shoots to climb

upward to the higher level of host trees.

These various climbing strategies not only form structural components in

any tropical forests and subtropical forests but also indicate different succession stages

of forests. Generally, tendril climbers are the early succession species whereas twiners

and root climbers are an indication of late succession species (DeWalt et al., 2000).

Several studies suggested that high proportions in species richness of twiners may

indicate that the forests may approach to a relatively late state of succession (Cai et al.,

2009; Chittibabu & Parthasarathy, 2001; DeWalt et al., 2000).

2.5 Liana climbing strategies and their limiting factors

The presence of suitability of host trees is an important factor influencing

liana access to the canopy. Most lianas climb their hosts using different climbing

strategies and require suitable structural supports to move upward. The studies in

several tropical and subtropical forests revealed that host trees characteristics may be

more important than environmental factors in determining the success of liana

colonization (Addo-Fordjour et al., 2009; van der Heijden & Phillips, 2008). Host

trees may differ in their suitability for liana colonization due to specific features such

as bark characteristics (Tally et al., 1996) or general factors such as trunk sizes


(Jiménez-castillo & Lusk, 2009; Nabe-Nielsen & Hall, 2002), trunk length

(Campanello et al., 2007). However, the suitability of support also varies with the

strategies by which lianas due to biomechanical constraints (Carrasco-Urra & Gianoli,

2009).

Host tree sizes related with tree age in that older trees, which are usually

large (Nabe-Nielsen & Hall, 2002), tend to be colonized by many liana stems (Pérez-

Salicrup & Meijere, 2005). Several studies have found more lianas associated with

larger trees. Addo-Fordjour (2008) suggested that trees with larger sizes supported

larger lianas than small host trees. The abundance of lianas per tree tended to increase

when increasing sizes of host trees (Campanello et al., 2007). Larger trunk diameters

were also less susceptible to liana colonization due to biomechanical limitations of

liana climbing abilities on larger trees (Putz & Chai, 1987). However, larger trees

carry more lianas than trees with smaller sizes (Nabe-Nielsen, 2001, Préses-Silcarb

2005). These might be because trees with larger trunk sizes might have experienced

longer exposure time to lianas infestation than tree small trees (Alvira et.al., 2004;

Préses-Silcarb, 2005).

It has been found that the limit of trunk sizes could vary with the types of

climbing strategies in tropical forests (Putz, 1984). For example, tendril climbers can

use smaller support than stem twiners (DeWalt et al., 2000). On the other hand,

adhesive root and hook climbers can use much larger supports (Cai et al., 2009).

Climbing mechanism often dictates which host trees lianas can climb (Pénalosa, 1982

but see Ludwig & Mainer, 2010). Host tree species may differ substantially in liana

load, especially among tropical canopy trees (Addo-Fordjour et al., 2009; Putz &

Chai, 1987).

Host tree bark texture has been reported to be important in determining the

presence of different climbing strategies (Chittibabu & Parthasarathy, 2001; Muñoz et

al., 2003). Smoothness of bark appears to be the feature of host trees with low liana

colonization or that are completely liana free (Campanello et al., 2007). Campanello et

al. (2007) suggested that host tree species with low liana load tended to have smooth

bark. For example, 70% of palm (species name) that had very smooth bark and large

leave are free of lianas (Campanello et al., 2007).



CHAPTER I

INTRODUCTION

1.1 Introduction

Lianas are woody climbing plants commonly found in most tropical forests

reaching the highest species diversity in tropical rain forests, evergreen forests, and

moist deciduous forests (Addo-Fordjour et al.,2009; Chittibabu & Parthasarathy, 2001,

Putz, 1984; Schnitzer & Bongers, 2002; Schnitzer, 2005; Senbeta et al., 2005).

However, several studies have suggested that environmental factors influence liana

distribution and abundance (DeWalt et al., 2010; Hu et al., 2010; Nabe-Nielsen & Hall

2002). Some studies have found that environmental factors may not directly influence

the liana distribution and abundance but rather the biotic factors such as the

availability of host trees and their characteristics could be related to liana abundance

(van der Heijden & Phillips, 2009; Nabe-Nielsen, 2001).

Because lianas rely on host trees for structural supports, a variety of

characteristics of host trees is important for the success of liana colonization. Most

lianas climb their hosts using different climbing strategies and require suitable

structural supports to move upward. Lianas employing the twiner and tendril strategies

appear to use hosts of small and medium host sizes (Nabe-Nielsen, 2001, Senbeta et

al., 2005), while lianas with adventitious roots and hooks require host trees that are

large enough to support their weight (Putz 1984). Moreover, host tree bark texture has

also been reported to be important in determining liana and host trees climbing success

(Muthuramkumar & Parthasarathy, 2001). Rough bark texture provides attachment

points for climbing better than trees with smooth bark texture (Campbell & Newbery,

1993; Muthuramkumar & Parthasarathy, 2001). Therefore, different characteristics of

host trees influence the presence of certain climbing strategy of lianas and may also be

related to liana diversity and abundance, which also influence forest dynamics.

Areas of biodiversity hot spot such as tropical forests in Thailand are also

rich in liana diversity. The Mo Singto Forest Dynamics Plot, which is a network of the

Natthida Khiewbanyang Introduction / 2

Center for Tropical Forest Science (CTFS), shows great liana diversity. The

relationship between liana and their limiting factors had previously been studied by

Lertpanich & Brockelman (2003). This study suggested that environmental factors did

not much determine the overall density and diversity of lianas in this plot. Host trees

with the diameter at breast height (dbh) less than 70 cm were highly colonized by

many liana ramets from different species. Therefore, biotic factors such as the

availability of host trees and their characteristics were probably related to liana

climbing success but these factors had yet been investigated. If the climbing strategies

of lianas were related to host tree characteristics, the following questions were

addressed.

1. What were the proportions of all liana climbing strategies in the Mo

Singto Forest Dynamics Plot?

2. Did different liana climbing strategies show selections for different

sizes and different bark textures of host trees?

1.2 Research Hypotheses

1.2.1 The diverse climbing strategies of lianas could be found at the Mo

Singto Forest Dynamics Plot and the proportion of lianas abundance with different

climbing strategies varied with types of supports and structures of the forests. It was

expected that twiner would be the dominant climbing strategy in the studied area

because twiners could generally use supports in the forest with high abundance of

small and medium trees.

1.2.2 Different liana climbing strategies were expected to select for

particular sizes and bark textures of host trees.

1.2.2.1 Twiners and tendril climbers were able to climb on

small sizes whereas lianas using other climbing strategies, such as the scramblers,

hook and adventitious root climbers could use larger host tree size.

1.2.2.2 Bark textures could also influence the presence of

lianas on host trees in that lianas preferred rough bark to smooth bark because liana


stem directly attach to host bark. Roughness of the bark promoted more areas that

lianas can climb host trees than smoothness of the bark.

1.3 Research Objectives

1.3.1 To record and classify climbing strategies of lianas in Mo Singto

Forest Dynamics Plot

1.3.2 To determine the proportions of different liana climbing strategies

1.3.3 To determine the relationships of different liana climbing strategies

on different sizes and bark textures of host trees


CHAPTER V

DISCUSSION

5.1 Lianas and host tree abundance

Liana diversity and abundance vary among sites throughout the tropical

forests (Putz, 1984; Schnitzer & Bongers, 2002; Schnitzer, 2012). Species richness of

lianas recorded in this study (123 species of 35 families found in 30.5 ha) was

relatively high when compared with some studies carried out in tropics. For instance,

Muthuramkumar & Parthasarathy (2000) recorded 75 species of lianas in a 30 ha plot

at Varagariar India, whereas Muoghalu & Okeesan (2005) recorded 49 species of

lianas in 20.5 ha plot at rainforest, Nigeria. Therefore, 123 liana species recorded in

this study is fairly high, considering the fact that only liana whose dbh of greater than

or equal to 3 cm were counted. Only the study conducted in the evergreen forest,

Southwest China, did identify the higher number of liana species (147 liana species in

15 ha plot) than the number obtained from this study (Cai, 2009).

Most liana ramets use some host trees such as S. sigun, N. melliferum, and

I. chevalieri, quite frequently. This could be due to the high density of these host tree

species in the study plot. Nevertheless, these trees only accounted for approximately

20% of all trees used by lianas. Thus, high abundance was not the prime choice for

structural supports. Other characteristics of host trees, such as trunk size and bark

texture, may influence variations of liana climbing strategies in forests as revealed by

several studies (Addo-Fordjour et al., 2009; Heijden et al., 2008; Muthuramkumar &

Parthasarathy, 2001; Nabe-Nielsen, 2001; Ludwig & Meiners, 2010).

5.2 Lianas climbing strategies

The Mo Singto Forest Dynamics Plot harbored high diversity of liana

species that exhibited various climbing strategies. The sampling liana ramets exhibited

five different climbing strategies with the twiner strategy as the most common

Natthida Khiewbanyang Discussion / 42

strategy. This finding was consistent with previous studies of tropical forests in that

the majority of liana species exhibited the twiner strategy (Chittibabu & Parthasarathy,

2001; DeWalt et al., 2000; Ghollasimood et al., 2012, Parthasarathy et al., 2004; Putz,

1984). For instance, in the Afromontane rain forests in Ethiopia, Senbeta et al. (2005)

reported that twiner was the dominant climbing strategy. This was similar to the

studies in evergreen forests in India (Parthasarathy et al., 2004), the dipterocarp forest

in Lambia National Park in Malaysia (Putz & Chai, 1987), tropical forests in China

(Cai et al., 2009), and the semi-deciduous rain forest in Ghana (Addo-Fordjour et al.,

2008).

However, some studies in subtropical forests in China reported that the

adventitious root strategy was the most common climbing method (Yuan et al., 2009)

while the tendril strategy was common in the dry evergreen forest (Parthasarathy et

al., 2004) and subtropical Atlantic forests (Campanello et al., 2007). Thus, climbing

strategies of liana species could vary in different areas. These diverse climbing

strategies could be a result of variations in liana species diversity. Although most liana

species usually exhibited one type of climbing strategies, they may use different

strategies in different regions. For instance, species in the genus Dalbergia found in

Afromontane rain forests in Ethiopia employed the twining strategy to climb host trees

(Senbeta et al., 2005). However, some species in this genus at the Mo Singto Plot

instead exhibited the tendril strategy. Another case in the coastal hill forest in

Malaysia reported that Tetracera indica employed the twiner strategy (Ghollasimood

et al., 2012), but this species at the Mo Singto Plot used the combination of twiner and

scrambler strategies. The fact that lianas used varieties of climbing strategies to climb

the forest canopy could be a result of adaptation (Putz, 1984; Senbeta et al., 2005).

It had been hypothesized that the twiner and tendril strategies were more

frequently found than the scrambler strategy in most tropical forests (Addo-Fordjour,

2008; Anbarashan & Parthasarathy, 2013; Cai 2009; Santos et al., 2009). Some studies

suggested that the scrambler strategy was not an effective method (Gentry 1991, Pual

& Yavitt, 2011; Schröder et al., 2013). The twiners tended to use leader shoots to

twine around their host trees and then used the replacing branches to climb upward if

they failed to find a support (Gerwing, 2004). However, twining ramets were

sometimes unable to maintain tensional forces and then fell of the host trees (Silk &


Holbrook, 2005). Putz (1984) suggested that slipping down the trunks was

occasionally observed. Based on the observation in this study, some liana ramets

frequently began their climbing by using the twiner strategy and then used the

scrambler strategy to climb on trellis of host trees to reach the canopy. The

combination of two strategies may help lianas to remain longer on the trunk of host

trees.

In addition, various climbing strategies not only form structural

components in any tropical forests and subtropical forests but also indicate different

succession stages of forest (Letcher & Chazdon, 2012). The different forms of

climbing habits might be related to differences in forest structure, dynamics, and age

(DeWalt et al., 2000, Putz & Chai, 1987). The tendril strategy was usually common in

forests with high density of small trees, which were characteristics of an early

succession stage while stem and branch twiners were be more common in later

successional forests (DeWalt et al., 2000). Twiners were able to climb small to

medium host tree trunks (Nabe-Nielsen, 2001; Senbeta et al., 2005). This was an

obvious advantage to twiners when the small supports were limited and climb more

successfully trees in the late succession stage of forest (Cai, 2009; Putz & Chai, 1987)

Based on this suggestion, the high proportion of liana ramets employing the twiner

strategy may indicate that the Mo Singto Forest may be reaching the late state of forest

succession. However, this assumption is tentative and needs to be tested.

Comparisons among tropical forest regions revealed that the lianas in this

seasonal evergreen forest exhibited as diverse climbing strategies as seen in other

regions. The dominant climbing strategy in an area may be caused in part by

differences in availability of suitable supports. Since the exhibition of liana climbing

strategies do not depend on species of host trees. It was noticed that liana ramets had

no preference for particular species of host trees because climbing strategies were a

result of adaptation.


5.3 Liana and host tree relationships

5.3.1 The relationship between liana ramets and host tree sizes

This study found that the relationships between lianas and their host tree

sizes were not random. The positive relationship indicated that small ramets tended to

use small hosts trees. As these ramets became larger, they also used larger hosts.

Although small ramets could use host trees of any sizes, they were found to climb

small trees more often than larger trees. It is possible that small trees were more

abundant in the area than larger trees. Some studies suggested that some of small

ramets were capable of growing under the low light environment and could only use

small host trees (Nabe-Nielsen, 2001). However, large ramets could not use hosts of

any sizes. They required certain sizes of trees that were able to support their sizes.

In this study, the size of trees being sampled was limited to the minimum

of 10 cm dbh. Nevertheless, liana ramets could potentially use hosts smaller than 10

cm dbh, which were not sampled in this study. Host tree seedlings were also expected

to be important for lianas in addition to small trees. Large host trees with dbh larger

than 70 cm could serve as supports for ramets to reach higher canopy and get direct

sunlight for rapid growth (Malizia & Grau, 2006). Areas with trees whose dbh smaller

than 70 cm usually had high liana density, but this correlation was not statistically

significant (Lertpanich & Brockelman, 2003). Some liana ramets could grow

independently without attaching themselves to host trees, and would climb on the

hosts when they were large. Upright individuals of some species could grow greater

than 2 m tall before requiring any supports that were large enough to support their

weight (Putz, 1984). Therefore, certain sizes of ramets require appropriate host tree

sizes for their successful colonization. Nevertheless, this pattern is not always

followed because some ramets of certain climbing strategies appear to be independent

of host tree sizes.

5.3.2 The relationships between liana climbing strategies and host tree

sizes

The twiner ramets appeared to be associated with host trees with diameter

< 50 cm dbh (Nabe-Nielsen, 2001; Senbeta et al., 2005). However, ramets using


twiners may be found on large host trees as reported in DeWalt et al. (2000) in that the

relative abundance of stem twiners in a tropical forest increased with forest stand age

(or trunk diameter). These twiners had attached to and grown on their hosts. As their

hosts became larger, the lianas also got larger.

The variation in climbing hosts of different sizes was observed for hook

climbers, but most of them usually used small host trees rather than larger host trees.

In contrast, Putz (1984) suggested that liana ramets employing the hook strategy could

use host trees regardless of their size. It was probable that some hook climbers only

required the presence of either branches or trunks of their hosts. As long as host trees

offered areas for attachment, they would be suitable for hook climbers. Nevertheless,

small branches usually provided better support than large trees because they offered

more surface areas for attachment. Thus, hook climbers tended to use small trees but

could also use trees of any sizes that provided areas for attachment.

Tendril ramets also used small host trees rather than other hosts of larger

sizes, consistent with the previous finding (Leicht-Young et al., 2010; Putz 1984). Due

to the fact that tendril climbers used sensitive organs to attach to their supports, large

host trees were likely too large for direct climbing because a tendril must wrap itself

around the supports. However, because the results from this study indicated that

tendril ramets were independent of host tree size, it would be similar to the hook

climbers that only required surface areas for holding on to the hosts.

A weak correlation between scrambler and host sizes possibly suggested

that these ramets only required structures for leaning. As a result, size of hosts may not

matter. This supported the suggestion that scramblers may depend on the density of

supports and climb more successfully trees in young forest where there are frequently

dense clusters of small trees (Campanello et al., 2007). Scramblers in this study were

found to climb not only their host trees but also other lianas. Similarly, the

combination of twiner and scrambler strategy also showed a weak correlation with

host tree sizes. Small host tree trunk was often used by twiners in the ascending state

but in the canopy scrambling ramets allowed the lianas to extend into the upper

canopy on branches of their host trees. The combination of twiner and scrambler

ramets would be more prevalent in younger forest and this trend was also observed in


this study. Therefore, small host trees provided better support for the combination of

twiner and scrambler ramets than less dense and large host trees.

Unlike other climbing strategies, the sizes of most adventitious root

climbers were medium and large. This strategy was found to use medium and large

host trees more often than other climbing strategies (Carrasco-Urra & Gianoli, 2009).

Because adventitious root climbers directly attached to trunks therefore, large host

trees provided more support area for attachment (Tally et al., 1996). Although this was

similar to the requirement of hook and tendril climbers, the adventitious root ramets

were present mostly in medium or large size and required bigger surface areas. Thus,

they were not independent of host tree sizes unlike other organ climbers. Since these

organ climbers appeared to be independent of host tree sizes but still required large

surface areas for attachment, other characteristics of host trees may be important.

5.3.3 The relationships between liana climbing strategies and different

host tree bark textures

Bark textures were related to climbing success with fewer lianas on trees

with smooth bark and more lianas on trees with rough bark that increased surface area.

It is likely that texture of bark help liana to climb successfully.

Although the result of this study revealed the frequent use of hosts with

slightly rough bark texture, it was as expected since most available host trees had

slightly rough bark texture. For instance, the highest percentage (8.66%) of lianas used

Sloenea sigun as hosts might be due to its high abundance and density in this plot.

Illex chavachirii with heavy lianas colonization also had slightly rough bark. These

species of host trees were the most abundant species in the area. Host trees with

smooth bark texture would be low in liana colonization because they lacked

attachment areas while rough and slightly rough bark offered more area. However, the

use of bark textures of lianas may depend on the climbing strategy.

Bark textures were likely to influence liana climbing success (Carse, 2000;

Carsten et al., 2002; Muñoz et al., 2003; Ladwig & Meiners; 2010). Although host

trees in the sampling area had slightly rough bark texture were used by ramets of

various climbing strategies. Some strategies did not often use this type of hosts. The


results from this study revealed that hook climbers tended to use host trees with the

slightly rough bark because this type of texture offered the attachment areas.

Not all climbing strategies required surface areas for colonization, some

strategies, such as the scramblers, only lean on the hosts with attachment. Host surface

areas as well as bark textures were not important.

Preference of particular bark texture of host trees was also obvious in

ramets using adventitious roots. These ramets showed their particular preference for

hosts with rough bark texture rather than size. Ramets of Rhus radicans used

adventitious roots to climb host trees were associated with trees with rough bark (Tally

et al., 1996). However, it has been suggested that host trees with strongly peeling bark

would be difficult for lianas to colonize using adventitious roots (Jiménez-Castillo &

Lusk, 2009). Rough bark was not stable surfaces for root attachment (Tally et al.

1996). For instance, Choerospondias axillaris, Michelia baillonii, Balakata baccata,

Prunus javanica, and Aphananthe cuspidata which had crack and fissure bark, had a

higher level of colonization by adventitious root ramets than Nauclea orientalis and

Alphonsea boniana, which had scale and vertical strips their bark. Despite this

observation, the roughness of the bark of host trees could promote lianas

establishment. Host trees with smooth bark may help deter liana colonization because

these host trees lack attachment sites.

The success of ramets of different climbing strategies may be explained by

differences in availability and suitability of physical supports. Comparisons among

tropical forest regions revealed that the lianas in this seasonal evergreen forest

exhibited diverse climbing strategies as seen in other regions. The study added to the

evidence that the presence and abundance of liana ramets with different climbing

strategies could be associated with size and bark texture of host trees not host species

in Mo Singto Forest. Host tree size was an important factor related to liana abundance,

as small host trees could facilitate liana colonization and provide a better support for

lianas. However, large host trees may become the necessary support by providing

canopy access.

The use of host trees for climbing supports also depend on the bark

textures of host trees. Rough bark of host trees was important characteristic

determining liana colonization. In addition, this unique morphology and habit of lianas


could explain for their distribution across forest succession but it is still unclear and

requires further investigation.

Natthida Khiewbanyang Results / 26

CHAPTER IV

RESULTS

4.1 Liana and host tree abundance

A total of 1,560 liana ramets examined in this study belonged to 52

common species distributing in 23 families. The ten most abundant liana species were

Uncaria scandens, Uncaria macrophylla, Tetracera indica, Premna flavescens,

Sphenodesme pentandra, Sabia limoniacea, Tetrastigma laoticum, Chonemorpha

fragrans, Caelospermum truncatum, Spatholobus harmandii (Table 4.1). The size of

liana ramets that reached host trees varied considerably ranging from 3 cm to 19 cm

dbh. The number of liana ramets per individual host tree ranged from 1-6 with average

of 1.27 ± 0.29 ramets.

These liana ramets attached to 1,251 host trees belonging to 120 species in

25 families (Appendix A). Of these 1,251 host trees, the top ten most abundant species

were Ilex chevalieri, Sloanea sigun, Mastixia pentandra, Symplocos cochinchinensis,

Dipterocarpus gracilis, Nephelium melliferum, Gironniera nervosa, Gonocaryum

lobbianum, Knema elegan, and Cinnamomum subavenium. Although liana ramets

appeared to show host tree species preference (2 = 3681.86, df = 119, p < 0.05), the

majority host tree species being used as structural supports were actually the most

abundant species in the study area (Table 4.2). For example, S. sigun had the highest

proportion (8.66%) of trees being held the lianas. Other host tree species also often

being used for supports were N. melliferum (7.77%), and I. chevalieri (5.91%).


Table 4.1 List of liana species and climbing strategies of 52 common species with dbh

≥ 3 cm in 30.5 ha plot. AD = Adventitious root; HO = Hook; SC = Scrambler; TD =

Tendril; TW = Twiner; TS = Twiner and Scrambler

Family Species nameNumber of

individuals in 30.5 ha plot

Density/haClimbing strategy

Rubiaceae Uncaria scandens 693 23.10 HO

Rubiaceae Uncaria macrophylla 436 14.53 HO

Dilleniaceae Tetracera indica 300 10.00 TS

Verbenaceae Premna flavescens 264 8.80 TS

Verbenaceae Sphenodesme pentandra 259 8.63 TS

Sabiaceae Sabia limoniacea 258 8.60 TW

Vitaceae Tetrastigma laoticum 219 7.30 TD

Apocynaceae Chonemorpha fragrans 181 6.03 TW

Rubiaceae Caelospermum truncatum 172 5.73 TW

Leguminosae Spatholobus harmandii 165 5.50 TW

Celastraceae Celastrus approximata 163 5.43 TW

Apocynaceae Ichnocarpus polyanthus 160 5.33 AD

Rubiaceae Morinda villosa 151 5.03 TW

Rubiaceae Morinda umbellata 137 4.57 TW

Rubiaceae Uncaria laevigata 131 4.37 HO

Convolvulaceae Neuropeltis racemosa 124 4.13 TW

Celastraceae Salacia chinensis 121 4.03 TS

Leguminosae Dalbergia horrida 119 3.97 TD

Convolvulaceae Erycibe elliptilimba 117 3.90 TWMenispermaceae Diploclisia glaucescens 116 3.87 TS

Melastomataceae Diplectria barbata 115 3.83 SC

Convolvulaceae Erycibe subspicata 110 3.67 TW

Apocynaceae Aganosma harmandiana 105 3.50 TS

Annonaceae Cyathostemma micrantha 98 3.27 TD

Hernandiaceae Illigera pierrei 98 3.27 SC

Apocynaceae Pottsia laxiflora 95 3.17 TW

Rubiaceae Oxyceros longiflora 94 3.13 HO

Vitaceae Tetrastigma aff.pyriforme 93 3.10 SC

Connaraceae Roureopsis stenopetala 91 3.03 SC

Annonaceae Desmos dumosus 86 2.87 SC

Annonaceae Fissistigma oblongum 84 2.80 TS

Rhamnaceae Ventilago denticulata 82 2.73 TS

Gnetaceae Gnetum montanum 79 2.63 TS

Dilleniaceae Tetracera loureiri 77 2.57 TW

Annonaceae Uvaria cordata 60 2.00 TD


Table 4.1 List of liana species and climbing strategies of 52 common species with dbh

≥ 3 cm in 30.5 ha plot. AD = Adventitious root; HO = Hook; SC = Scrambler; TD =

Tendril; TW = Twiner; TW, SC = Twiner and Scrambler (Cont.)

Family Species nameNumber of

individuals in 30.5 ha plot

Density/haClimbing strategy

Annonaceae Uvaria lurida 55 1.83 SC

Moraceae Ficus sagittata 54 1.80 TW

Apocynaceae Urceola micrantha 54 1.80 AD

Elaeagnaceae Elaeagnus conferta 53 1.77 SC

Menispermaceae Hypserpa nitida 51 1.70 TS

Euphorbiaceae Phyllanthus reticulatus 50 1.67 SC

Connaraceae Rourea minor 49 1.63 TS

Combretaceae Combretum latifolium 48 1.60 TW

Thymelaeaceae Linostoma pauciflorum 47 1.57 TD

Apocynaceae Melodinus fusiformis 44 1.47 TW

Urticaceae Poikilospermum suaveolens 43 1.43 SC

Leguminosae Mucuna macrocarpa 40 1.33 TW

Piperaceae Piper ribesoides 39 1.30 TS

Piperaceae Piper retrofractum 36 1.20 TD

Leguminosae Dalbergia velutina 36 1.20 SC

Gnetaceae Gnetum macrostachyum 33 1.10 TS

Apocynaceae Strophanthus caudatus 32 1.07 TS


Table 4.2 The top ten most abundance species of host trees ≥ 10 cm dbh in the Mo

Singto Forest Dynamics Plot, KhaoYai National Park, Thailand.

4.2 Liana climbing strategies

From all lianas sample recorded, the most common climbing strategy was

the twiner strategy, which was found in 480 liana ramets from 16 liana species

followed by the scrambler strategy seen in 330 liana ramets from 10 species. The

tendril strategy, which was the third most common strategy, was found in 150 liana

ramets from six species while the hook strategy, which was less common, was present

in 120 liana ramets from four species. The least common strategy in the study area was

the adventitious root found in 60 ramets from two species (Table 4.3). Most liana

species exhibited one type of climbing strategies (2 = 544.615, df = 5, p < 0.05). For

example, all species in the family Convolvulaceae used the twiner strategy to climb up

host tree canopy. Most species of lianas (57.14%) in the family Apocynaceae only

used the twiner strategy. In the subfamily Papilionoideae (family Leguminosae), a few

species, all ramets in the genus Dalbergia, used the tendril strategy to climb host trees.

All species in the genus Uncaria used the hook strategy. However, there was an

exception in that some liana species employed the combination of twiner and

scrambler strategies. The liana species that employed this climbing combination

included Aganosma harmandiana, Diploclisia glaucescens, Fissistigma oblongum,

Gnetum macrostachyum, Gnetum montanum, Hypserpa nitida, Piper ribesoides,

Tree speciesNumber of individuals

Number of host tree climbed by lianas

% Liana climbing

Ilex chevalieri 1021 73 5.83

Sloanea sigun 1008 107 8.55

Mastixia pentandra 768 56 4.48

Symplocos cochinchinensis 768 55 4.39

Dipterocarpus gracilis 757 56 4.48

Nephelium melliferum 735 96 7.67

Gironniera nervosa 708 50 4.05

Gonocaryum lobbianum 590 41 3.32

Knema elegan 479 39 3.16

Cinnamomum subavenium 474 35 2.83


Premna flavescens, Rourea minor, Salacia chinensis, Sphenodesme pentandra,

Strophanthus caudatus, Tetracera indica, and Ventilago denticulata.

Table 4.3 The number of liana species and the proportion of individual ramets with

various climbing strategies in 30.5 ha seasonal evergreen forest, Mo Singto Forest

Dynamics Plot.

Climbing strategyNumber of individual

rametsProportion (%)

Number of species

Proportion (%)

Single climbing strategy

Twiner 480 30.77 16 30.77

Scrambler 330 21.15 10 19.23

Tendril 150 9.61 6 11.54

Hook 120 7.70 4 7.69

Adventitious root

Combination of two strategies

60 3.85 2 3.85

Twiner and Scrambler 420 26.92 14 26.92

Total 1560 100 52 100


4.3 Liana and host tree relationships

4.3.1 Relationships between liana ramet sizes and host tree sizes

Host trees and ramet sizes appeared to be correlate (r2 = 0.166, p < 0.05)

(Figure 4.1). Host trees with larger dbh tended to support liana ramets with larger dbh.

Small host trees could support small ramets. Medium host trees were found to be

related with medium size of ramets. Large ramets tended to use medium and large host

trees. The reverse was not true because large host trees could support all sizes of liana

ramets as small ramets less than 6 cm dbh were present on some of the large host trees.

Although a general trend showed the positive correlation between host

trees and liana ramet sizes, different size group of host trees did not equally provide

structural supports for the ramets. The majority of the lianas used small host trees

particularly of 20-30 cm dbh size. The abundance of liana ramets slightly decreased

when increasing host tree diameter and distinctively decreased at the medium host tree

size class (40-70 cm dbh).

Some ramets did not show any correlation with host tree sizes but rather

host tree bark texture. Host trees with slightly rough bark were usually used by liana

ramets of every size (Figure 4.2). However, patterns of some relationships were

detected under certain conditions, especially for different climbing strategies.


Figure 4.1 The relationship between liana and host tree sizes with a regression line

and r2 = 0.166, p < 0.05.

Figure 4.2 The percentage of liana ramets on three types of bark textures (smooth.

slightly rough and rough), for each of three size classes.


4.3.2 Relationships between liana climbing strategies and host tree

sizes

There was a positive correlation between liana climbing strategies and host

tree sizes. The results showed that the number of ramets using different climbing

strategies appeared to use certain sizes of host trees.

Ramets employing the twiner strategy showed a strong positive correlation

with host tree sizes. The sizes of these twiners increased with increasing host tree size

(r2 = 0.457, p < 0.05) (Figure 4.3). Most twiners were small and mostly use small host

(Figure 4.9a). Small twiners could twine around small host trees. Both medium and

large twiners were usually found on medium host trees and some of medium and large

twiners could also use large host trees. However, the abundance of twining ramets

decreased with increasing size of host trees. Observed values in each class of host tree

sizes showed that the twiners appeared to use certain sizes of host trees (2 = 15.27, df

= 2, p < 0.05). Although small host trees were the most abundant size of host trees, the

twiners showed more frequent use of small hosts than usual.

The sizes of hook climbers showed a weak correlation with host tree sizes

(r2 = 0.175, p < 0.05) (Figure 4.4). The majority of liana ramets using this climbing

strategy were small. These small ramets tended to use small host trees. However, small

ramets were found to use medium and large host trees as well. The abundance of liana

ramets in each class of host tree sizes showed that the hook climbers were found to use

small host trees but they also used hosts of other sizes as available (Figure 4.9b).

There was a weak positive correlation between ramets using the tendril

strategy and host tree size (r2 = 0.208, p < 0.05) (Figure 4.5). The majority of these

tendril climbers were also small and usually found on small host trees (Figure 4.9c).

Medium ramets appeared to use medium host trees as well as large host trees.

However, the data showed that in fact these ramets were independent of host tree sizes

(2=2.926, df = 2, p<0.05).

The correlation between size of ramets employing the scrambler strategy

and size of host trees also had a weak correlation (r2 = 0.069, p<0.05) (Figure 4.6).

Most of these ramets were small size. Although most of them were found on small

host trees, they appeared to be independent of host tree sizes (Figure 4.10d).


The similar situation was found in ramets that used the combination of

twiner and scrambler strategies to climb host trees. The size of these ramets was

weakly dependent on host tree sizes (r2 = 0.066, df = 2, p < 0.05) (Figure 4.7).

Although most of these ramets were small, they were able to attach to wide range of

host sizes (Figure 4.10e).

A weak correlation between ramets using the adventitious root strategy

and host tree sizes was found (r2 = 0.121, df = 2, p < 0.05) (Figure 4.8). Unlike other

ramets of other climbing strategies, ramets with this climbing strategy were mostly

medium in size. The majority of these ramets tended to use medium host trees (Figure

4.10f). Observed values in each class of host tree sizes showed that the adventitious

root climbers appeared to use medium size hosts more often than expected (2=15.27,

df = 2, p < 0.05).


Figure 4.3 The relationship between liana size and host tree size of the twiner strategy

with a regression line and r2 = 0.457, p < 0.05.

Figure 4.4 The relationship between liana size and host tree size of the hook strategy



Figure 4.5 The relationship between liana size and host tree size of the tendril strategy


Figure 4.6 The relationship between lianas diameter and host tree size of the

scrambler strategy with significant regression line at r2 = 0.069, p < 0.05.


Figure 4.7 The relationship between liana size and host tree size of the combination of

twiner and scrambler strategy with a regression line and r2 = 0.066, p < 0.05.

Figure 4.8 The relationship between liana size and host tree size of the adventitious

root strategy with significant regression line at r2 = 0.121, p < 0.05.


Figure 4.9 The host tree diameter size ≥10 cm dbh used by liana ramets of ≥3 cm dbh

at Mo Singto Forest Dynamics plot, Khao Yai National Park Thailand: (a) ramets

showing the twiner strategy, (b) the hook strategy, and (c) the tendril strategy.

(a)

(b)

(c)


Figure 4.10 The host tree diameter size ≥10 cm dbh used by liana ramets of ≥3 cm

dbh at Mo Singto Forest Dynamics plot, Khao Yai National Park Thailand: (d) ramets

showing the scrambler strategy, (e) the combination of twiner and scrambler strategy,

and (f) the adventitious root strategy.

(d)

(e)

(f)


4.3.3 Relationships between liana climbing strategies and different host tree bark

textures

There were proportional differences between the three bark types (smooth,

slightly rough and rough) used by lianas. Ramets regardless of their climbing

strategies tended to use hosts with slightly rough bark rather than rough or smooth

bark (Table 4.4). Ramets mainly used host trees with slightly rough bark (61.54%) and

less frequently used other bark types; only 32.24% of hosts with rough bark and 6.22%

of hosts with smooth bark.

Liana ramets that used different climbing strategies were found to be

correlated with particular types of bark texture of host trees (Table 4.4). Host trees

with slightly rough bark were being used with the highest percentage by the number of

individual ramets. However, the 2 test verified that hook climbers tended to use host

trees with slightly rough bark of host trees more often than expected (2=11.52, df = 2,

p < 0.05). On the other hand, adventitious root climbers required host trees with rough

bark (2= 8.32, df = 2, p < 0.05). In contrast, ramets employing the other climbing

strategies were independent of host tree bark textures.

Table 4.4 Percentage of individual ramets and the percentage of individual ramets for

each climbing strategy belonging to different bark textures of host trees. AD =

Adventitious root; HO = Hook; SC = Scrambler; TD = Tendril; TW = Twiner; TS =

Twiner and Scrambler

Percentage of individual ramets for each

climbing strategyHost tree bark

texture

Percentage of

individual rametsTW SC TD HO AD TS

Smooth 6.22 1.85 1.73 0.44 0.12 0.51 1.54

Slightly rough 61.54 19.29 13.71 5.25 6.64 1.73 15.90

Rough 32.24 9.61 5.70 3.91 1.92 1.60 9.49

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LIANA CLIMBING STRATEGIES AND RELATIONSHIP WITH HOST …