The Effects of Human Impacts on
Cave and Karst Biodiversity:
Thailand Component
Final Report
Project code:
RE-THL-005
Final project report for the period: February 2002—February 2004
Reporting date:
25 April 2004
Project start:
February 2002
Project completion:
May 2004
Total donor budget:
≈ 60 000
Forest
Insect
Group
The Effects of Human Impacts on Cave and Karst Biodiversity: Thailand Component
is funded by the ASEAN Regional Centre for Biodiversity Conservation and the European
Commission
“This document has been produced with the financial assistance of the ASEAN Regional Centre for Biodiversity Conservation (ARCBC) funded by the European
Commission. The views expressed herein are those of the author/authors and can therefore in no way be taken to reflect the official opinion of the ARCBC and/or the
European Commission nor those of the Department of National Parks and Wildlife/Royal Forest Department (Thailand)”
The cover art shows three important cave animals encountered during this study. From top to
bottom: an Amblypygid or Whip-spider - large and important predators; a Rhaphidophorid “cave
cricket” - very abundant scavengers and Schistura magnifluvis (Kotellat) a loach type of fish.
Cover artwork courtesy Piyawan Niyomwan (Amblypygid), Wichai Sorapongpaisan (Rhaphidophorid),
Sansanee Amornpurinant (S. magnifluvis) and Sopitpang Luanghiran (S. magnifluvis).
Cover layout by Dean Smart, Parkphoom Sangklin and Robert Cunningham.
“Beauty and the beast”: The beautiful tower karst through which Tham Jet Khot passes,
with a tarmac car park suitable for 10 or more tour buses in the foreground. At the start of
field sampling for this project in January 2003 the road to this undisturbed end of Tham
Jet Khot was a narrow dirt track. By our final sampling trip in May 2003 there was a
tarmac road and this car park. The frightening pace at which this development has taken
place highlights the urgent need for more research to understand the impacts of tourists on
caves.
NOTICE
This document is the final report for the project “The Effects of Human Impacts on Cave
and Karst Biodiversity: Thailand Component”. It had been prepared in accordance with
the ARCBC recommended format, as detailed in Annex VI of the project contract, however
some modifications have been made to better accommodate flow and formatting.
v
Preface
Nearly 4 years ago I accepted an invitation from Dean Smart, a speleologist who has
been conducting research into the caves of Thailand since 1993, to lead this project.
There were two reasons behind my acceptance. Firstly, I am an entomologist and a
conservationist, and talks with Dean led me to realise the important of the biodiversity
of caves and their vulnerability to disturbance. In Thailand, like elsewhere, the biggest
problem for proper management of cave biodiversity is the lack of data thus making
research an urgent priority. This project is our small effort to address this problem.
Secondly, the involvement of Robert Cunningham in the project gave me great confidence. Robert is keen on research experimental design and data analysis and his input
was crucial for the success of the project.
The project is now complete and I hope the benefits will be clear to all. In addition to scientific results, with clear management implications, the project has created
new collaborations and friendships between institutions in Thailand and Indonesia. In
Thailand this project is the first ever to bring ichthyologists, speleologists, arachnologists and entomologists to work in an integrated way on cave fauna. The project is
also the first in Thailand to establish a good collection of cave terrestrial arthropods.
Speaking personally I’ve discovered just how fascinating and magnificent caves are, and
I know I am not alone in this discovery. I know that my own personal discovery will
remain with me for the future and hope that the other benefits of the project are also
perpetuated.
Of course, without the support of the ARCBC and EU no project would have been
possible. We are very much indebted to both for their financial and other support
which has allowed us to shed a little light on cave biodiversity in South-east Asia.
I would like to thank personally all those people who have helped this project,
especially my project staff who worked diligently, overcoming many obstacles, and
supported me throughout the duration of the project.
Finally, I would like to thank Dean Smart who introduced me to caves and their
importance, and brought the project idea to us.
Chaweewan Hutacharern
Project Leader
July 2004
i
ii
Contents
Contents
iii
List of Tables
vii
List of Figures
ix
1 Frontmatter
1.1 Project participants . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1.2 List of acronyms used . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1.3 Executive summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
1
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Project Administration and Budget
2 Project administration
2.1 Current workplan status . . . . . .
2.1.1 ASEAN and EU cooperation
2.1.2 Cave biodiversity assessment
2.1.3 Report results . . . . . . . .
2.1.4 Management . . . . . . . .
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3 Budget
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II
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Technical reporting
4 Introduction/Project background
4.1 Caves and disturbance by human visitors . . . .
4.2 Not all caves are equally sensitive to disturbance
4.3 The need for evidence . . . . . . . . . . . . . .
4.4 South-east Asian caves and their biodiversity . .
4.5 What biodiversity? . . . . . . . . . . . . . . . .
4.6 Our principal goal . . . . . . . . . . . . . . . . .
5 Project objectives
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6 Materials and methods
6.1 Experimental approach and design . . . . . . . . . . . . . . .
6.1.1 Human disturbance — the primary experimental factor
6.1.2 Secondary factors . . . . . . . . . . . . . . . . . . . . .
6.1.3 Co-factors . . . . . . . . . . . . . . . . . . . . . . . . .
6.2 Study area and study caves . . . . . . . . . . . . . . . . . . .
6.2.1 Study area . . . . . . . . . . . . . . . . . . . . . . . . .
6.2.2 Study caves . . . . . . . . . . . . . . . . . . . . . . . .
6.3 Disturbance assessment . . . . . . . . . . . . . . . . . . . . . .
6.4 Biodiversity assessment . . . . . . . . . . . . . . . . . . . . . .
6.4.1 Terrestrial Arthropods . . . . . . . . . . . . . . . . . .
6.4.2 Fish . . . . . . . . . . . . . . . . . . . . . . . . . . . .
6.5 Physical and environmental parameters . . . . . . . . . . . . .
6.6 Data management and analysis . . . . . . . . . . . . . . . . .
7 Results
7.1 Disturbance . . . . . . . . . . . . . . . . . . . . . . .
7.2 Terrestrial Arthropods . . . . . . . . . . . . . . . . .
7.2.1 Catch summary and sampling efficiency . . . .
7.2.2 Abundance . . . . . . . . . . . . . . . . . . .
7.2.3 Species richness . . . . . . . . . . . . . . . . .
7.2.4 Biodiversity indices . . . . . . . . . . . . . . .
7.2.5 Relationship between biodiversity parameters
scores . . . . . . . . . . . . . . . . . . . . . .
7.2.6 Indicator species analysis . . . . . . . . . . . .
7.2.7 Community composition . . . . . . . . . . . .
7.2.8 Morphological modifications for hypogean life
7.3 Fish . . . . . . . . . . . . . . . . . . . . . . . . . . .
7.3.1 Catch summary and sampling efficiency . . . .
7.3.2 Abundance . . . . . . . . . . . . . . . . . . .
7.3.3 Species richness . . . . . . . . . . . . . . . . .
7.3.4 Biodiversity indices . . . . . . . . . . . . . . .
7.3.5 Relationship between biodiversity parameters
scores . . . . . . . . . . . . . . . . . . . . . .
7.3.6 Indicator species analysis . . . . . . . . . . . .
7.3.7 Community composition . . . . . . . . . . . .
7.3.8 Condition index changes . . . . . . . . . . . .
7.3.9 Morphological modifications for hypogean life
7.4 Physical and environmental data . . . . . . . . . . .
8 General discussion
8.1 The parts . . . . . . . . . . .
8.1.1 Sampling effectiveness
8.1.2 Catch summary . . . .
8.1.3 Disturbance effects . .
8.1.4 Season effects . . . . .
8.1.5 Distance effects . . . .
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and disturbance
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and disturbance
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9 Conclusions and recommendations
9.1 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
9.2 Management recommendations . . . . . . . . . . . . . . . . . . . . . .
9.3 Research recommendations . . . . . . . . . . . . . . . . . . . . . . . . .
77
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10 Acknowledgements
79
III
81
8.2
8.1.6 Indicator species . . . . . . . . . . . . . . . .
8.1.7 Community composition . . . . . . . . . . . .
8.1.8 Morphological modifications for hypogean life
8.1.9 Environment and physical . . . . . . . . . . .
8.1.10 General fauna . . . . . . . . . . . . . . . . . .
8.1.11 Limitations . . . . . . . . . . . . . . . . . . .
The whole . . . . . . . . . . . . . . . . . . . . . . . .
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Appendices
A Publications and Presentations
A.1 Technical and administrative reports . . . . . . . . . . . . . . . . . . .
A.2 Presentations at scientific meetings . . . . . . . . . . . . . . . . . . . .
A.2.1 Workshop on “The effects of human impacts on cave and karst
biodiversity” . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
A.2.2 ARCBC Regional Research Grant Conference: Building bridges
between ASEAN and EU researches[sic] . . . . . . . . . . . . .
83
83
84
B Taxonomic checklist
87
C Location, cave and station maps and sketches
C.1 Chumphon . . . . . . . . . . . . . . . . . . . . .
C.1.1 Tham Than Nam Lot Yai . . . . . . . .
C.2 Phangnga (Amphur Mueang) . . . . . . . . . .
C.2.1 Tham Phung Chang . . . . . . . . . . .
C.2.2 Tham Tapan . . . . . . . . . . . . . . .
C.3 Phangnga (Amphur Thap Phut) . . . . . . . . .
C.3.1 Tham Thong . . . . . . . . . . . . . . .
C.3.2 Tham Nam 1 . . . . . . . . . . . . . . .
C.4 Satun . . . . . . . . . . . . . . . . . . . . . . .
C.4.1 Tham Jet Khot . . . . . . . . . . . . . .
C.4.2 Tham Khong Kha Lot . . . . . . . . . .
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84
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D Contact sheet
119
Bibliography
123
v
vi
List of Tables
2.1
2.2
Workplan activity progress at the end of project . . . . . . . . . . . . .
Cooperation with visiting international speleological expeditions . . . .
10
11
6.1
6.2
Climatic summary of the study area . . . . . . . . . . . . . . .
Table of study cave names, codes and cave ends with greatest
disturbance . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Details of disturbance scoring levels . . . . . . . . . . . . . . .
28
6.3
7.1
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relative
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29
31
ANOVA table of effects of disturbance, season and distance on TA abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of disturbance, season and distance on TA observed species count . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of disturbance, season and distance on TA estimated species richness . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of disturbance and season on TA Fisher’s α . .
ANOVA table of effects of disturbance, season and distance on TA H0 .
Summary of TA ISA scores and their significance for disturbance . . . .
Summary of TA ISA scores and their significance for distance . . . . . .
ANOVA table of effects of disturbance, season and distance on fish abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of inflow-outflow, season and distance on fish
abundance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of disturbance, season and distance on fish observed species count . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of disturbance, season and distance on fish estimated species richness . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of disturbance, season and distance on fish
Fisher’s α . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
ANOVA table of effects of disturbance, season and distance on fish H0 .
Summary of Fish ISA scores and their significance for disturbance . . .
Summary of Fish ISA scores and their significance for distance . . . . .
Summary of ANOVA for the condition index (k) and body length of ten
of the more common fish species . . . . . . . . . . . . . . . . . . . . . .
Environmental and physical characteristics of the seven study caves . .
63
64
B.1 Checklist of fish species collected during the study . . . . . . . . . . . .
88
7.2
7.3
7.4
7.5
7.6
7.7
7.8
7.9
7.10
7.11
7.12
7.13
7.14
7.15
7.16
7.17
vii
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viii
List of Figures
6.1
6.2
7.1
7.2
7.3
7.4
7.5
7.6
7.7
7.8
7.9
7.10
7.11
7.12
7.13
7.14
7.15
7.16
7.17
7.18
7.19
7.20
C.1
C.2
C.3
C.4
C.5
C.6
C.7
C.8
C.9
Experimental layout schematic showing paired disturbance levels and
sampling stations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Map of southern Thailand showing the locations of the study caves . .
Disturbance scores for each of the seven study caves . . . . . . . . . . .
Randomised species accumulation, singleton and doubleton curves. . . .
Overall TA abundance for each of the seven study caves . . . . . . . . .
TA abundance by disturbance level, season and distance . . . . . . . .
TA observed species count by disturbance level, season and distance . .
TA estimated species by disturbance level, season and distance . . . . .
TA Fisher’s α by disturbance level and season . . . . . . . . . . . . . .
TA H0 by disturbance level, season and distance . . . . . . . . . . . . .
TA CCA ordination plot showing the 14 study cave ends . . . . . . . .
Mosaic plot of TA Order level taxonomic composition differences between disturbance levels . . . . . . . . . . . . . . . . . . . . . . . . . .
Randomised species accumulation, singleton and doubleton curves. . . .
Overall fish abundance for each of the seven study caves . . . . . . . .
Fish abundance by disturbance level, season and distance . . . . . . . .
Fish abundance by inflow-outflow, season and distance . . . . . . . . .
Fish abundance by disturbance level, season and distance . . . . . . . .
Fish estimated species by disturbance level and distance . . . . . . . . .
Fish Fisher’s α by disturbance level and distance . . . . . . . . . . . . .
Fish H0 by disturbance level, season and distance . . . . . . . . . . . .
Fish CCA ordination plot showing the 14 study cave ends . . . . . . .
Changes in Systomus binotatus condition index with season . . . . . .
26
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Location of Tham Than Nam Lot Yai in amphur Sawi, Chumphon . . . 93
Tham Than Nam Lot Yai floor plan . . . . . . . . . . . . . . . . . . . . 94
Sketches of sampling stations in Tham Than Nam Lot Yai . . . . . . . 95
Locations of Tham Phung Chang and Tham Tapan in amphur Mueang,
Phangnga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Tham Phung Chang floor plan . . . . . . . . . . . . . . . . . . . . . . . 98
Sketches of sampling stations in Tham Phung Chang . . . . . . . . . . 99
Tham Tapan floor plan . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sketches of sampling stations in Tham Tapan . . . . . . . . . . . . . . 102
Location of Tham Thong and Tham Nam 1 in amphur Thap Phut,
Phangnga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
ix
C.10 Tham Thong floor plan . . . . . . . . . . . . . . . . . . . . . . . . . . .
C.11 Sketches of sampling stations in Tham Thong . . . . . . . . . . . . . .
C.12 Tham Nam 1 floor plan . . . . . . . . . . . . . . . . . . . . . . . . . .
C.13 Sketches of sampling stations in Tham Nam 1 . . . . . . . . . . . . . .
C.14 Location of Tham Jet Khot, amphur La-ngu, Satun province, and Tham
Khong Kha Lot, amphur Pa Bon, Phattalung province . . . . . . . . .
C.15 Tham Jet Khot floor plan . . . . . . . . . . . . . . . . . . . . . . . . .
C.16 Sketches of sampling stations in Tham Jet Khot . . . . . . . . . . . . .
C.17 Tham Khong Kha Lot . . . . . . . . . . . . . . . . . . . . . . . . . . .
C.18 Sketches of sampling stations in Tham Khong Kha Lot . . . . . . . . .
C.19 Legend for station maps . . . . . . . . . . . . . . . . . . . . . . . . . .
105
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D.1 A selection of project photographs — set 1 . . . . . . . . . . . . . . . . 120
D.2 A selection of project photographs — set 2 . . . . . . . . . . . . . . . . 121
x
Chapter 1
Frontmatter
1.1
Project participants
The project has had eleven active participants:
1. Ophart Chamason
2. Robert Cunningham
3. Prasong Hemapan
4. Chaweewan Hutacharern
5. Sommai Janekitkarn
6. Ruenchit Phukthair
7. Watana Sakchoowong
8. Patpimon Sawai
9. Dean Smart
10. Patchanee Vichitbandha
11. Chavalit Vidthayanon [terminated October 2002]
1
2
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
List of acronyms used
A list of acronyms as applied to the names of the caves in the study is provided in
table 6.2 (page 29), other acronyms used are listed here.
ARCBC ASEAN Regional Centre for Biodiversity Conservation
ASEAN Association of South East Asian Nations
ATBI All Taxon Biological Inventory
CCA Canonical Correspondence Analysis
DNPW Department of National Parks, Wildlife and Plant Conservation, Thailand
EU European Union
FIG Forest Insect Group
IDH Intermediate Disturbance Hypothesis
ISA Indicator Species Analysis
KU Kasetsart University
NRCT National Research Council of Thailand
PAR Photosynthetically Active Radiation
RTU Recognisable Taxonomic Unit
TA Terrestrial Arthropods
2
3
1.3
Executive summary
Project Administration and Budget
The project has been completed on time (February 2002–February 2004), within budget
(see summary table below and chapter 3) and without any major difficulties. All
planned activities (chapter 2, § 2.1) and major objectives (see below and chapters 4
and 5) have been met and the project has produced scientifically interesting results
(see below). The project has been a success.
A list of the projects objectives is provided below:
1. Determine the effects of human impacts on cave biodiversity
2. Investigate spatial, ecological and trophic distribution patterns for cave biota
3. Make recommendations for the management of cave biodiversity
4. Create cave biota reference collections in a suitable museum
5. Integrate data into a project database
6. Disseminate results
7. Bring together Indonesian and Thai researchers in a joint project working towards
improving local and regional cave and karst management
8. Create working links between Association of South East Asian Nations (ASEAN)
and European Union (EU)
Budget
heading
Human Resources
Travel + Perdiem
Equipment
Supplies
Other
Contingency
Total
Annual budget ( )
Year 1
Year 2
16 120
16 120
6 540
3 448
4 000
0
5 000
5 900
400
450
1 011
1 011
33 071
26 929
Total
budget
32 240
9 988
4 000
10 900
850
2 022
60 000
3
Total
expenditure
36 378.25
11 519.21
4 717.67
5 957.25
1 364.45
59 936.83
Total expenditure as
% of total budget
112.84
115.33
117.94
54.65
160.52
99.89
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Technical results
Introduction There are more people visiting caves now than at any time in history
and the trend is for further increases. Cave ecosystems are widely regarded as being
particularly sensitive to disturbance. The belief that cave ecosystems are particularly
sensitive is based largely on anecdotal and fragmentary evidence with very little quantifiable data. These three facts make it imperative that research be conducted to
demonstrate with quantifiable data the effects of human disturbance on cave ecosystems — particularly on their biodiversity. In this study we investigated the effects
of human disturbance on the biodiversity of Terrestrial Arthropods (TA) and fish in
high-energy stream caves in south-east Asia. High energy caves are often regarded as
being less susceptible to disturbance than low-energy caves, but again there is a paucity
of solid data.
Materials and methods The effects of human impacts on cave biodiversity were
investigated by conducting a “natural experiment”with seven through-caves in southern
Thailand. Each cave had both a relatively disturbed and a relatively undisturbed end,
these ends being two treatments blocked within the seven caves which were treated as
replicates. In addition to the effects of disturbance the effects of season (period of the
dry season) and distance from cave entrance were investigated (see chapter 6, § 6.1 and
§ 6.2). Terrestrial arthropods and fish were selected as target taxa by which to assess
biodiversity (chapter 4, § 4.5).
Terrestrial arthropod sampling was conducted at six stations (each 50 m apart),
while fish sampling took place twice at three stations (upstream and downstream of
TA stations 1, 3 and 5). Sampling took place on three occasions: early dry season, the
middle of the dry season and late in the dry season. Terrestrial arthropods were sampled
by quantified hand collecting from three habitat strata: floors, interfaces and walls. All
TA were collected and preserved and later identified to recognisable taxonomic units
in the laboratory. Fish were sampled from quantified search areas by searching and
collecting with a scoop net and electro-fishing equipment. Fish were identified, weighed,
measured and, in most cases, released in the field.
In order to confirm our a priori assignment of disturbed and undisturbed cave ends
a disturbance artefact based scoring system was developed and used to quantify disturbance.
The effects of disturbance and other experimental factors on biodiversity was examined by assessing the effects on abundance, species richness, biodiversity measures
and community composition. In the case of fish the condition index of fish was also
investigated.
Results Scoring of caves for their disturbance score confirmed the validity of our a
priori relative disturbance assignments.
A total of 4 079 TA representing 519 RTUs1 from 25 orders were collected. One
thousand six hundred and eighty one fish specimens were collected representing 57
species, 40 genera from 17 families. No obviously troglobitic TA or fish were collected.
1
Recognisable Taxonomic Units as the majority of material is unidentified and probably unidentifiable in the immediate future
4
5
The cave (replicate block) effect was significant for both TA and fish for almost
all the biodiversity parameters examined. The significance of the effect confirming the
effectiveness of the blocked design.
Both TA and fish show clear effects of human disturbance on biodiversity with
both taxa showing highly significant differences in abundance, species richness (observed species count and estimated species) and α biodiversity indices (Fisher’s α and
H0 ) between disturbance levels. Disturbed cave ends showing consistently greater abundance, richness and diversity than undisturbed ends.
With respect to season, TA also showed season effects with the mid-dry season
having lower abundance, species richness and α diversity than other seasons. Fish
showed no significant seasonal effects with respect to any of the biodiversity parameters
but one species, Systomus binotatus, showed changes in condition index according to
season and distinct patterns for disturbed and undisturbed cave ends.
The distance effect was contrary to season, being clear in fish but absent in TA. Fish
showed a clear decline in abundance, species richness and α diversity with increasing
distance from the cave entrance. Terrestrial arthropods showed no clear pattern.
Neither TA nor fish had any species that could be identified by indicator species
analysis as indicators of undisturbed cave ends, though there were three TA and two
fish species identifiable as indicators of disturbance.
Terrestrial arthropods also had two species that were significant indicators of stations furthest from cave entrances, with a further nine species indicators of cave entrances. Fish had seven species that were indicators of cave entrances and none for the
inner parts of caves.
With regard to community composition, both TA and fish showed the greatest similarities within caves (regardless of disturbance level), far overwhelming any effects of
disturbance. Among TA the Araneae and Orthoptera (dominated by Rhaphidophoridae) were relatively over-represented in undisturbed cave ends while the Coleoptera
were over-represented in disturbed cave ends.
Thirty eight potential confounding physical and environmental factors were examined and none were found to show any significant disturbance level differences and no
factor or combination of factors were found to be adequate predictors of disturbance.
Conclusions and Recommendations The results of this study show clearly that disturbance due to human visitation to stream caves causes increases in the abundance,
species richness and α biodiversity of both TA and fish. This implies that the overall
biodiversity of a stream cave changes when subjected to such disturbance. The belief
that ’high energy’ stream caves are less susceptible to disturbance than other types of
cave may need to be re-evaluated. This study also discovered several species of TA and
fish that were indicators of disturbance and their presence in a cave demonstrates that
biodiversity impacts have already reached the levels seen here. Management of stream
caves for tourism needs to be more protective of the cave through control of visitors
and applying strategies to mitigate impacts. Out of this study and meetings between
cave and karst researchers came several recommendations concerning further research.
These centre on expanding our knowledge of taxonomy and ecology of caves in ASEAN
and the mechanisms by which impacts on biodiversity occur.
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Key words: Thailand, cave biodiversity, cave management, fish, terrestrial
arthropods, tropical caves
6
Part I
Project Administration and Budget
7
Chapter 2
Project administration
There ain’t no rules around here.
We’re trying to accomplish
something.
Thomas Edison
2.1
Current workplan status
The project is effectively complete with almost all the proposed workplan activities
completed (Table 2.1).
Further details on the activities in each of the four major workplan headings are
provided below.1
2.1.1
ASEAN and EU cooperation
ASEAN cooperation
During April 4–11 2003 six of the “The Effects of Human Impacts on Cave and Karst
Biodiversity: Thailand Component” project team (R. Cunningham, S. Janekitkarn,
R. Phukthair, W. Sakchoowong, P. Sawai, D. Smart) visited Indonesia as part of an
exchange of visits between this project and our Indonesian sister project (“The Effects
of Human Impacts on Cave and Karst Biodiversity: Indonesian Component”). Our
visit included discussions with Indonesian team members at Bogor, examination of
project materials and facilities at the Museum Zoologicum Bogoriense at Cibinong,
and a four day visit to the caves of the Maros karst on Sulawesi where we visited some
of the caves studied by the Indonesian project. The trip went well and was interesting,
productive and enjoyed by all involved.
During May 25–28 2003 three of the “The Effects of Human Impacts on Cave and
Karst Biodiversity: Indonesian Component” project team (Ristiyanti Marwoto, Cahyo
Rahmadi and Yayuk Suhardjono) visited Thailand as part of the exchange of visits.
1
the details provided are for the period February 2003–February 2004, periods prior to this having
been reported in earlier progress reports to ASEAN Regional Centre for Biodiversity Conservation
(ARCBC)
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Table 2.1: Workplan activity progress at the end of project
Workplan
activity
ASEAN and EU cooperation:
a) Exchange trip Maros-Thailand
b) Exchange trip Thailand-Maros
Cave biodiversity assessment:
a) purchase caving equipment and books
b) purchase collecting equipment
c) purchase computer
d) preliminary survey trips
e) 1st collecting trip
f) 2nd collecting trip
g) 3rd collecting trip
h) sorting and curation of specimens
i) create reference collections
j) update databases
k) collate results
Report results:
a) Semi-annual report year 1
b) Annual report, year 1
c) Final report
Project management:
% Completion at
end of project
100%
100%
100%
100%
100%
100%
100%
100%
100%
95%
95%
90%
100%
100%
100%
100%
100%
During their time in Thailand the Indonesian team attended the workshop “The effects
of human impacts on cave and karst biodiversity” (see section A.2.1, page 84) and
visited four of the Thai study caves and the coastal karst landscapes of Phangnga bay.
During the ARCBC “Regional Research Grant Conference: Building bridges between
ASEAN and EU biodiversity researches[sic]” several team members (R. Cunningham,
C. Hutacharern, D. Smart), as well as Y. Suhardjono from our Indonesia sister project,
along with other ASEAN scientists participated in a special focus group, discussing
ideas for future ASEAN-wide cooperation on cave and karst research.
The workshop“Sampling standardization on cave and karst fauna”originally planned
(by the Indonesian team) for April 2003 in Indonesia did not receive funding from ARCBC and so has yet to take place. We hope there will be the opportunity to participate
in this workshop in the future.
EU and other international cooperation
Dean Smart (United Kingdom) and Robert Cunningham (Australia) have been actively
involved throughout all phases of the project and Dean Smart is expected to continue
speleological research in Thailand.
During the processing of specimens we have been assisted by several EU and other
international taxonomists:
Martin Ole Odderskov (Denmark)
10
Chapter 2. Project administration
11
Kazuo Ogata (Japan)
Sadahiro Ohmomo (Japan)
Jan Peterson (Denmark)
Peter Weygoldt (Germany)
We expect that further material from the project will also be examined by interested
researchers in the future.
Project staff have collaborated with several visiting foreign speleological expeditions, see details in Table 2.2.
Table 2.2: Cooperation with visiting international speleological expeditions
Group
Date
Societe Speleo Ariege Pays D’Olmes (FR)
July 2003
Shepton Mallet Caving Club (UK)
December 2003
University of Bristol Speleological Society (UK) December 2003
2.1.2
Cave biodiversity assessment
Collecting and sorting of specimens, and collation and analysis of results has been
completed and is reported on in Part II of this report.
Some workplan activities in this section are not 100% complete, a brief description
of their status is provided below.
Sorting and curation of specimens Specimen sorting is 100% complete as are most
curation activities, the remaining needs for curation are for the specimen vials to
be properly bulked, labeled and stored.
Create reference collections Working reference collections (used during sorting)
are complete but some of these materials could be developed into more permanent
reference collections.
Update databases All cave databases have been updated. Biodiversity data is also
maintained in statistical databases as well as working files in Microsoft Excel
and/or ASCII format. However, the fish data seems suitable for development
into a publicly accessible database of taxa and this should be done at some time
in the future.
2.1.3
Report results
With the submission of this report all reporting requirements for the project have been
fulfilled, further details of reporting can be found in Appendix A (page 83). In addition
to materials published and presented to date it is hoped that material from the project
will be suitable for publication in the future.
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
2.1.4
Management
Management proceeded more-or-less in accordance with our expectations and the workplan — the project being carried out to a successful conclusion.
12
Chapter 3
Budget
When money speaks, the truth
remains quiet.
Russian proverb
Financial reporting details (forms RG 01-02, RG 02-02, RG 03-02 and RG 04-02)
have been submitted earlier with the closing of ARCBC accounts on January 18, 2004
and these details are not repeated here. The only budget information provided in this
report is the budget summary table included in the Executive Summary (page 3).
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
14
Part II
Technical reporting
15
Chapter 4
Introduction/Project background
a tenuous balance that teeters on
the brink of doom
a description of cave ecosystems
McReynolds 1975
4.1
Caves and disturbance by human visitors
Despite the fact that we consider the “caveman” period long past the reality is that
there are more people (cavemen and cavewomen) visiting more caves now than at any
other time in human history. Watson et al. (1997) gave an estimate of 20 million cave
visitors to more than 650 lit“show caves”), and with increasing interest in adventure and
ecotourism, these numbers seem likely to continue rising. Like most human activities,
visiting caves is a disturbance sensu latu to the cave with consequent impacts on the
cave ecosystem.
Disturbances to caves by visitors range from the sublime e.g. changes in cave
atmosphere (Ferández-Cortés et al., 2003) or damage to hypogean root communities
by the introduction of energy from epigean sources (English and Blyth, 2000) to the
ridiculous, e.g. use of explosives in the search for a train of “lost Japanese gold” in
Thailand — “treasure-based” tourism ((The Bangkok Post, 2001)
Unfortunately, as result of their typically inherently relatively steady-state, cave
ecosystems are considered to be among the most sensitive of ecosystems such that even
minor disturbances may have severe adverse1 impacts.
The sensitivity of cave ecosystems and their increasing exposure to disturbance by
human visitation, and other factors, are resulting in caves becoming increasingly endangered ecosystems (Spate and Hamilton-Smith, 1991). Activities of humans visiting
caves induce greater threat than “the activities of quarry operators and other users, or
abusers, of karst areas”, and they are not alone in this view (Howarth, 1983).
1
adverse here implying any change from their existing state
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
4.2
Not all caves are equally sensitive to disturbance
Of course, as in all of Nature, there is variation and not all cave ecosystems are equally
sensitive to disturbance. Cave managers/researchers
often distinguish a dichotomy between “low-energy” and “high-energy” caves. Lowenergy caves having relatively low energy inputs (in terms of both kinetic and chemical
energy) when compared to high-energy caves. A paradigm has become established
that low-energy caves are especially sensitive to disturbance while high-energy caves
are more robust (Vermeulen and Whitten, 1999; Watson et al., 1997). This paradigm
often forms the basis of decisions related to cave management and zoning 2 with highenergy caves being regarded as more suitable for visitation (perhaps as show caves) than
low-energy caves because of their perceived robustness. Unfortunately speleothems
associated with low-energy caves are often more spectacular and/or delicate and so
more visit-worthy than those that are able to form in high-energy caves (where high
kinetic energy processes make delicate speleothem formation difficult), while at the
same time some high-energy caves are unsuitable for visitation because of the dangers
posed by their high-energy state, flash flooding of stream caves being a good example
— thus providing a quandary for managers.
While this paradigm is well established and we do not doubt its application to abiotic features such as speleothems, sedimentary or archaeological deposits, etc., we are
not convinced that the same applies to biological components of cave ecosystems. There
are many ecosystems that could be characterised as high-energy, certainly much higher
energy than even high energy cave systems, which are nevertheless sensitive to disturbance, e.g. epigean riparian or intertidal ecosystems. Even Spate and Hamilton-Smith
(1991) in their discussion of energy levels and disturbance on biological communities
concede that what might be more important is whether the disturbance “produce[s]
changes considerably outside the normal parameters”. This view is more in accord with
our thinking that energy levels are not the most important determinant of susceptibility of a cave to disturbance, rather aspects of disturbance itself (such as intensity,
frequency, predictability) relative to the normal range encountered by the cave are more
important. We do acknowledge the value of the high-energy/low-energy dichotomy as
a useful rule of thumb for management as there is often a correlation between energy
state and the range of normal parameters, but feel that more care should be exercised
in it’s use.
4.3
The need for evidence
The real problem is lack of convincing evidence. We share the view of Sutherland (2000)
on the need evidence-based conservation and the evidence with respect for disturbance
effects on caves, and for differences between caves of different energy levels — especially
cave biodiversity — is sparse at best. The little evidence that does exist is largely
anecdotal and fragmentary.
There are certainly many anecdotal reports about human disturbance through visitation affecting cave ecosystems but few studies are well documented, and even fewer
2
when such noble matters as management and zoning are considered at all!
18
Chapter 4. Introduction/Project background
19
have sound quantitative information. Spate and Hamilton-Smith (1991) recounts many
stories of human impacts on cave biodiversity but remarkably few, if any, of these are
based upon quantitative data. Even recent research titles as promising as “Natural and
human impacts on invertebrate communities in Brazilian caves” (Ferreira and Horta,
2001) being at best semi-quantitative and poorly analysed
In a quite extensive review of impacts on caves by Clarke (1997) the impacts of
visitors were largely ignored. The comments of Howarth (1983) calling for more work
to be done on the effects of human visitation to caves are as applicable today as they
were twenty years ago.
One of the principle goals of this current work is to provide sound quantitative
evidence showing the effects of human disturbance on cave biodiversity.
4.4
South-east Asian caves and their biodiversity
In this study we examined the impacts of human disturbance on biodiversity not in
low-energy temperate zone caves but in higher-energy tropical caves — in particularly
those in SE Asia.
Caves and cave biodiversity in south-east Asia is poorly known compared to many
other regions of the world, particularly Europe and North America. The cave fauna of
the entire region remains virtually unknown and the situation for species biology and
ecology is even worse (Deharveng and Bedos, 2000). This is unfortunate because the
few caves that have been studied, for example: Batu Caves, Malaysia (McClure, 1965);
the caves of Mulu, Sarawak (Chapman, 1982); Tham Chiang Dao, Thailand and the
Ngalau Surat–Gua Salukkan Kallang system and Batu Lubang, Indonesia (Deharveng
and Bedos, 2000), all demonstrate the existence of a highly diverse and important fauna
including many troglobitic and relictual species. Gua Salukkan Kallang being recognised as the only subterranean biodiversity “hotspot” in all of Asia (Culver and Sket,
2000). This lack of knowledge prevents proper assessment of the biological significance
and vulnerability of caves in ASEAN and severely hampers the development of improved cave management. We expect that our work will be a small contribution to the
current state of knowledge.
4.5
What biodiversity?
Biodiversity is a rather broad concept including diversity at the genome, species and
ecosystem levels (Anonymous, 1998). Obviously when we consider the effects of human
disturbance on caves we should consider biodiversity at all levels. Equally obviously,
this is almost never possible. In our case we have chosen to look at species level biodiversity. Even at this level the scale of the problem is enormous given the tremendous
level of species biodiversity across a wide range of taxa. While we agree, at a cerebral level, with Janzen (1997) that an All Taxon Biological Inventory (ATBI) provides
many benefits we, at a more visceral level, also agree with (New, 1998) that ATBIs “are
usually not feasible” and more simplified approaches often yield useful results (New,
1999). Of the two common alternatives to ATBIs for assessing biodiversity impactsmuseum type collecting (a wide range of taxa but typically not well quantified) and
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
indicator taxa (well quantified but restricted to a narrow range of taxa) we find little satisfaction. Rather we find the “shopping basket” approach of Stork (1995) more
compelling, this approach being to sample quantitatively from a rather broad range of
taxa, though certainly not comparable to the scope of an ATBI.
In this study we chose terrestrial arthropods and fish3 as our “shopping items”. As
we were interested in the impact of human disturbance on both terrestrial and aquatic
habitats these two taxa provided the coverage we were looking for. Some further reasons
for our selection of these taxa are provided below.
Why terrestrial arthropods? Arthropods are simply the most diverse of all living
organisms with estimates of species richness ranging from 5 million to as many as 100
million species (Ødegaard, 2000). As Platnick (1991) so clearly puts it,
“Speaking about biodiversity is essentially equivalent to speaking about
arthropods. In terms of species, other animal and plant groups are just
gloss on the arthropod scheme.”
Caves, are no different from the world at large and arthropods are also the most
diverse group of organisms in caves (Welbourn, 1999).
In addition to their diversity arthropods are also amongst the most abundant of life
forms, Stork (1988) estimating that the number of arthropods in one hectare of rain
forest exceeds 42 million.
As well as their hyper-diversity and great abundance (and in part as a result of it)
arthropods (and other invertebrates) have major functional roles in most ecosystem
processes and as Wilson (1987) describes it “they are the little things that run the
world”.
In addition to their importance, arthropods and invertebrates in general have many
practical aspects that make them suitable for biodiversity assessment and monitoring
(see Basset et al. (1998) and references therein).
Why Fish? We chose fish for an assortment of reasons. Fish are by far the most
diverse of the vertebrates, with more than 25 000 species (Nelson, 1994)
Some species are recognised and used as indicators of water quality (Barbour et al.,
1999) Thailand is particularly rich in troglobitic cave fish with ≈ 10% of the global total
of known cave fish species (Helfman et al., 1997; Vidthayanon et al., 1997). Fish are
also particularly vulnerable to the effects of human visitation for fishing is a common
reason for local people to visit caves.
4.6
Our principal goal
As outlined above, while disturbance by humans visiting caves is becoming an increasingly important issue, quantitative data is almost non-existent. The main goal of this
project was to address this problem; in particular, to answer the following question:
Does disturbance caused by human visitation to high-energy cave ecosystems in SE Asia, have an impact on the biodiversity of such caves??
3
principally Osteichthyes
20
Chapter 4. Introduction/Project background
21
Addressing this question would involve a systematic survey where quantifiable data
would be collected and then subjected to testable, statistical analyses.
In addition to this main goal, the project had a variety of ancillary objectives that
are discussed briefly in the following chapter.
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
22
Chapter 5
Project objectives
In addition to the primary objective introduced in the preceding chapter, the project
had seven other secondary objectives as detailed in our proposal and these are listed
here:
1. Investigate spatial, ecological and trophic distribution patterns for cave
biota: in addition to the effects of disturbance, we also looked at variation among
season, distance into the caves, taxa differences etc.
2. Make recommendations for the management of cave biodiversity: the
results of this work will likely have direct implications for future management and
research of high-energy tropical caves and recommendations will be given based
on those results.
3. Create cave biota reference collections in a suitable museum: as part
of the processing of specimens, voucher specimen and other reference collections
will be established and maintained.
4. Integrate data into a project database: much of the project data has been
entered into and maintained in databases.
5. Disseminate results: as the results of this work will be of value to cave managers and fellow researchers, the results should be disseminated to interested
parties.
6. Bring together Indonesian and Thai researchers in a joint project working towards improving local and regional cave and karst management:
an important part of the project was to foster collaboration between Thai and
Indonesian researchers with the hope that partnerships will be developed to maintain research effort in the future.
7. Create working links between ASEAN and EU: in addition to links within
ASEAN, the development of links to EU researchers was an important part of the
project.
23
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
24
Chapter 6
Materials and methods
No one would now dream of testing
the response to a treatment by
comparing two plots, one treated
and the other untreated.
Fisher and Wishart 1930
6.1
Experimental approach and design
As made clear in the introduction, the most important goal of this project was to
provide quantitative, experimental data on the effects of human disturbance on cave
biodiversity — examining TA and fish as focal groups.
The ideal experiment would have been a manipulative one where we started with
several replicate undisturbed caves, or ends of caves, and randomly assigned some
to be disturbed while others are left undisturbed as controls (with the possibility of
temporal monitoring to examine the dynamics of disturbance). However, as might
be expected, this ideal situation was not available. As is often the case with any
larger-scale ecological experiments, such as this one, we needed to adopt a “natural”
experiment where we examined and tested quantitatively the effects associated with
“treatments” that had already been “assigned”.
Despite the limitations imposed by being a “natural experiment” we have adopted
a rigorous experimental design with the aim of producing sound, quantified data. The
details of the experimental design are described below.
6.1.1
Human disturbance — the primary experimental factor
Disturbance levels
The primary experimental factor is human disturbance, herein referred to as disturbance. Disturbance has two levels: relatively disturbed and relatively undisturbed,
herein referred to as disturbed and undisturbed .
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Fish sampling area
TA sampling area
Entrance
1
2 3 4
Stream
5
Unsampled
buffer zone
Entrance
6
5 4
3 2
1
6
Disturbed end
Undisturbed end
Figure 6.1: Experimental layout schematic showing paired disturbance levels and
sampling stations. Inset shows the detail of a sampling station, showing TA and fish sampling areas in relation to a station mid-point
Replicate blocks
With respect to disturbance, and several other factors (both secondary such as described
below and ancillary, such as day on which sampling took place), the seven caves included
in the study are treated as replicate blocks, each cave having both a disturbed and
an undisturbed end (Figure 6.1). In addition to the spatial pairing with each of the
treatments in the one cave we also sampled both ends of the cave simultaneously
(on the same day and starting at the same time each day (1000 hs)) thus providing
temporal blocking. Blocking techniques such as this allows us good statistical control
of “unwanted” inter-cave variation (uniqueness being a well known phenomena of caves
as well as allowing simple straightforward contrasts with less restrictive assumptions
than other designs (Box et al., 1978; Sokal and Rohlf, 1995).
6.1.2
Secondary factors
In addition to disturbance there were several secondary factors which were either of
interest to us directly or which we wished to control as part of the experimental design.
Secondary factors of direct interest to us were season and distance into the cave from
the entrance, herein distance.
Season Because of our interest in energy availability in caves following the wet season
flood we sampled three times during the dry season1 , as detailed below.
Trip 1: nominally the beginning of the dry season, after the last wet season
flood when we expected that energy, in the form of flood debris, would be at a
1
wet season sampling is very dangerous as the caves are prone to severe flooding, see Figure D.1(c),
page 120
26
Chapter 6. Materials and methods
27
maximum (although that energy may not be readily available to other than early
stage decomposers at this stage)
Trip 2: nominally the middle of the dry season
Trip 3: nominally the end of the dry season when we expect energy to be at its
lowest. While this is nominally the end of the dry season predicting the end of
the dry season is of course difficult and at least two caves (Tham Khong Kha Lot
and Tham Than Nam Lot Yai) had flooded just prior to our final sampling trip
Distance Each cave end had six sampling stations with each sampling station’s midpoint being located at 50 m intervals, with the first station starting at the cave entrance
(mid-point 10 m from the entrance) and the last ending at 270 m (mid-point 260 m) into
the cave. As most of the chosen caves exceed 700 m in length this allowed a minimum
of more than 100 m buffer between each disturbance level (Figure 6.1). Because one
cave (Tham Nam 1) was shorter than the surveyed length2 it had only five stations
from each end and the final stations of each end were only separated by an interval of
10 m. Terrestrial arthropods were sampled at every station while fish were only sampled at stations 1, 3 and 5. Further details of the nature of the stations with respect
to terrestrial arthropods and fish are provided in sections 6.4.1 and 6.4.2 respectively.
Water flow In addition to these secondary factors, of direct interest to us we were
also concerned about the direction of the flow of water through the caves, especially as
some aspects of disturbance, particularly water-borne disturbance has the potential to
“flow” along with the water. This would be especially important for fish. As a result
of this concern we also tried to block and balance so that the number of caves with
disturbed inflows and disturbed outflows was similar, as similar as possible given an
uneven number of caves.
6.1.3
Co-factors
In addition to the primary and secondary experimental factors, we measured a wide
range of other parameters associated with the study caves. Among these were the
scoring of disturbance (to validate our a priori assignment of disturbance levels) and
the measurement of a wide range of physical aspects of the caves and a wide range of
environmental parameters. Details of these measurements and parameters are provided
below (sections 6.3 and 6.5).
6.2
6.2.1
Study area and study caves
Study area
Southern Thailand lies north of the equator between latitudes 6 ◦ and 11◦ and from
longitude 098◦ to 102◦ E. It is bounded on the west by the Andaman Sea and to the
east by the Gulf of Thailand. The area lies within the Sundaic biogeographical region
2
which was only discovered on the first sampling trip
27
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
and possesses a faunal and floral assemblage with closer affinities to Malaysia, Borneo,
Java and Sumatra than to the rest of Thailand. The northern boundary of the Sundaic
region passes through the narrowest part of peninsula Thailand at about 11 ◦ north.
Geologically, southern Thailand consists of two mountain chains running northsouth with a broad area of low elevation in between and coastal plains on the outsides.
The western chain is formed mainly of Permo-Carboniferous sedimentary rocks with
later granite intrusions. Granite forms the bulk of the eastern chain, in which the
highest mountain of southern Thailand is located - Khao Luang, 1 835 m. Karst rocks
are widespread throughout the area occurring as outliers on the flanks of the mountains.
There are two main karst-forming rocks: the Ordovician (dark, thinly bedded limestone
restricted to the south and east) and the Permian (pure, massive limestone mostly in
the north and west). Karst and cave development is more prominent in the Permian,
which forms the spectacular tower karst of Phangnga and Krabi. The Ordovician is
more subdued with shorter, rounded landforms (Kiernan (1988) and references therein).
The total number of known caves in the area exceeds 500 (Dunkley (1995); unpublished
data). However, exploration and surveying continues to find previously undocumented
caves (Gray, 2001; Smart and Cunningham, 2001, 2002) and the actual total is probably
several times this number. The longest caves are all active stream caves collecting
drainage off adjacent, non-carbonate rocks and passing through the bases of towers
and ridges, etc.
The climate of the area is influenced by two distinct monsoon seasons, westerly
(May to November) and easterly (November to January). The prevailing monsoons
tend to dump their rain on the first coastline they encounter. Consequently, the west
coast receives much of its rain during the westerly monsoon and vice-versa for the east
coast. Mean annual climatic data for locations near to the study caves are as follows
(data from Climatology Division (1990)):
Table 6.1: Climatic summary of the study area
Meterological
station
Chumphon
Phuket Airport
Satun
a
b
Study area
Chumphon
Phangnga
Satun
Rain
(mm)
2 000
2 500
2 200
Rainy seasona
May - Nov
Apr - Nov
Apr - Nov
Monsoon
direction
West
West
West
Temperatureb
(◦ C)
26.7
27.4
27.5
Relativeb
Humidity (%)
82
80
78
the rainy season is defined as the period when average monthly rainfall exceeds 150 mm
mean annual average
6.2.2
Study caves
Seven through-caves were selected (after considerable initial survey work documented
in Smart and Cunningham (2001, 2002)) in which there was a clear apparent relative
difference in disturbance between ends, this being the basis on which our initial a priori
assignment of disturbance levels was made. All caves were located in three provinces
in southern Thailand with further location details in Table 6.2 and Figure 6.2.
All caves had a perennial water flow (required for fish sampling) and a main stream
28
Chapter 6. Materials and methods
29
passage longer than 700 m3 . In addition to these criteria caves were selected to allow
for blocking of water flow effects as described earlier (section 6.1.2).
Table 6.2: Table of study cave names, codes and cave ends with greatest relative
disturbance
Cavea
Tham
Tham
Tham
Tham
Tham
Tham
Tham
Code
Tapan
TPN
Jet Khot
JKT
Khong Kha Lot
KKL
Phung Chang
PCH
Thong
TNG
Than Nam Lot Yai NLY
Nam 1
NM1
Area
Phangnga
Satun
Satun
Phangnga
Phangnga
Chumphon
Phangnga
Disturbed end
Outflow
Outflow
Inflow
Outflow
Inflow
Inflow
Inflow
a
the caves are ordered from greatest relative difference in disturbance between ends (Tham Tapan)
to least relative difference (Tham Nam 1), as explained further in Figure 7.1
3
note however the problem of Tham Nam 1 referred to earlier
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
12
o
Chumphon
1. Tham Than Nam Lot Yai
10
1
*
o
North
4
3
2
08
**
**
5
o
Phangnga
Limestone karst
2. Tham Phung Chang
3. Tham Tapan
4. Tham Thong/Lot
5. Tham Nam 1
*6 * 7
Satun
Km
o
100
098
Ó DS 2003
o
100
o
0
102
06
6. Tham Jet Khot
7. Tham Khong Kha Lot
o
Figure 6.2: Map of southern Thailand showing the locations of the study caves
30
Chapter 6. Materials and methods
6.3
31
Disturbance assessment
In order to test our a priori assessment of relative disturbance differences between cave
ends we devised and used a system to score human disturbance. This was based on the
assumption that disturbance is associated with physical artefacts that we are able to
measure. The system was not intended to provide an absolute measure of disturbance
(although it may with modification) rather it was intended to show relative differences
between ends and among caves. To make the assessment seven types of disturbance
artefacts were assessed and assigned to one of four states with a score of 1-4 (Table 6.3).
Scoring was carried out at every station and was conducted independently by two people
(DS and RC), one per team, during trips two and three. The scores for each of the
disturbance artefacts were then summed to produce a disturbance score for each station.
In addition to scoring disturbance artefacts within caves we also assessed accessibility
to each cave entrance, again using a scoring system of 1-4 (1-jungle bash, 2-footpath;
3-road or track nearby; 4-car park). These accessibility scores were added to each
station at the respective cave end. The final disturbance score for each station being
the sum of the seven disturbance artefacts and the accessibility score. In addition to
station scores cave ends and whole caves were rated according to the mean disturbance
scores of their stations.
Table 6.3: Details of disturbance scoring levels
Artefact type
Litter
Footprints
Graffiti
Lighting
Speleothem damage
Development
Other
1
None
None
None
None
None
None
None
2
3
4
1 piece
2 + same
2 + multiple
Few
Footpath
Brainy floor
1 piece
2 + same
2 + multiple
1 type
2
3+
1 broken 2 - 5
6+
1 simple 2 + simple Complex
1 type
2-5
6+
Notes:
Litter: 2 + same = several pieces of litter of the same type that could have been
dropped by one group, e.g. sweet wrappers. 2 + multiple = several pieces
probably dropped by different groups or at different times, e.g. cardboard
incense box and plastic water bottle.
Footprints: Footpath was assessed to be a clear walking track. Brainy floor
was where a sediment floor had been completely compacted. The footprint
scores were adjusted for stations where the floor was completely flooded or
comprised only bare rock. The adjustment looked at the score of two “end”
stations where a sediment floor was present and assumed that the stations
in between would have similar scores.
Graffiti: 2 + same = several pieces of graffiti of the same type that seems likely
to have been written by 1 group, e.g. all in red paint. 2 + multiple =
several pieces probably written by different groups or at different times,
e.g. different media or dates.
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Lighting: evidence of light sources, e.g. discarded batteries, spent calcium carbide, candle wax, burnt bamboo, electric wiring and lights.
Speleothem damage: the number of broken stalactites and stalagmites, or the
amount of trampling damage on flowstone.
Development: the presence and number of any infrastructure items for visitation, e.g. wooden plank bridges (simple), steps cut into floor (simple),
concrete pathway (complex), bridges (complex), boat (complex).
Other: simple totaling of instances of physical impacts not listed above, e.g.
fires, burning incense, catching fish or bats, feeding fish, digging up the
floor, a dam, smoking, dropping food, etc.
6.4
Biodiversity assessment
Specimen collection was carried out by two teams working simultaneously in each end of
the same cave. Each team consisted of four people - two people for collecting terrestrial
arthropods and two for fish.
6.4.1
Terrestrial Arthropods
Field methods
Quantified sampling for TA was conducted by hand collecting 4,5 in three habitat strata
viz. floors, walls and interfaces (the interface between the floor and wall) and at 12
stations in each cave (six stations in each end Figure 6.1). Sampling was stratified in
order to provide more complete coverage across the range of major cave habitats and
to allow comparisons among habitats. This later aim was not achieved as clumping
across habitats was required in order to have reliable data for analysis and so the
habitat factor is not considered further in this report.
Each of the three habitat strata had its own particular sampling methods, the
details being described below.
Floor At each station, six quadrats (0.5 m × 0.5 m) were randomly placed within the
floor area (being 20 m in length) of each station, three in the left and three in
the right. Two collectors spent 3 minutes searching (with the aid of a normal
electric caving light6 and using a count-down stop-watch) and hand collected all
TA encountered within the area of each quadrat. Searching included both an
initial surface search followed by disturbance of rocks, litter and the soil surface
in each quadrat.
4
we had hoped to use more collector-independent methods pitfall traps and Berlese/Tullgren
funnels but like others (i.e. Scharff et al. (2003)) found logistic difficulties precluded this
5
hand collecting means that the sample could be better described as a sample of macro-arthropods
(those being visible to the naked eye) and certainly the micro-arthropods community was not included
by this sampling method
6
initially we tried collecting under red-light as this disturbs and attracts arthropods less than
white-light but this was abandoned as several collectors found searching under red-light “too difficult”
32
Chapter 6. Materials and methods
33
Interface The interface is the zone where the cave floor and wall meet (over the same
area of the station as described for the floor). For the purposes of our sampling
it included ≈ 0.5 m of floor and ≈ 0.5 m of wall at the floor-wall intersection. In
order to avoid repeated collection of the same area of interface (as a result of
returning to the same station on three occasions) each 20 m section of interface
habitat (left and right walls) was divided into three parts of 6 m each (allowing a
1 m buffer between parts). On each sampling trip one part from each side, which
had not yet been sampled, was randomly chosen for sampling, thus ensuring
that no part was collected twice.7 Furthermore, adjacent left and right interfaces
and interface and wall habitats on the same side were never sampled on the
same occasion. These blocks were further divided into two 3 m units that were
sampled separately. Hand collecting was done by searching the interface and
all TA encountered were collected,8 there was no time limit and collecting was
complete once the interface had been traversed once.
Wall Wall sampling was essentially the same as that described for interface sampling.
The area of the wall sampled was that between ≈ 0.5 m and ≈ 1.5 m high on each
wall.
Samples from each sampling event (quadrat or part of transect) were kept stored
separately in polythene vials containing 80% ethanol along with a label indicating cave,
cave end, station, habitat type, sub-sample details, collector and date.
On some occasions and at some stations sampling was not possible because the
habitat did not properly exist (a flooded floor or interface), or it was too dangerous to
access (having to collect from a wall while swimming in a flooded stream). In these
cases these samples were regarded as having no terrestrial arthropods.
On one occasion it was not possible to collect the designated sample on account of an
inconveniently resident King Cobra (Ophiophagus hannah (Cantor 1836)). Strictly this
should be treated as “missing data”, however, we decided to regard it as another case
of “no terrestrial arthropods” as to introduce missing data would have unnecessarily
complicated the analysis.
Specimen sorting and identification
As is typically the case with tropical terrestrial arthropods it is not possible and in
our case, not necessary or desirable to spend the time identifying specimens to species
and so we adopted the morphospecies concept as recommended by Oliver and Beattie (1993). Despite criticisms (Goldstein, 1997), this seems to be the only practical
approach for the present (see also (New, 1998)). Here we refer to morphospecies as
Recognizable Taxonomic Units (RTU).
7
we did not take the same precautions with the quadrat sampling as typically the area sampled on
each occasion was only a small proportion of the total area available and so the chances of a randomly
placed quadrat falling in the same place were small. Furthermore the disturbed nature of the floor
(flooding, visitors, etc) made it much less likely to have sessile animals than the wall, sessile animals
being obviously much more subject to over-collecting
8
although some of the faster Amblypygi, Heteropodidae and Rhaphidophoridae did escape on
some occasions
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Specimens were initially sorted to Order and transferred to glass vials with fresh
80% ethanol. Orders were then assigned to six of us (RC, CH, WS, PS, DS, and PV)
for sorting to RTU, each sorter being responsible for all the specimens in one or more
Orders, this being a measure to help maintain consistency of RTUs.
Although most material has been sorted to RTU, some taxa have been determined
to Family, Genus or Species. Resources used to assist sorting include:
“Isopoda” (Brusca, 2003)
“Walking with woodlice: Identification key”The Natural History Museum (2003a,b)
“Arthropoda, Diplopoda, Polydesmida” (Lana, 2003)
“Centipedes and Millipedes-Myriapoda” (Australian Museum, 2003a)
“Class Diplopoda: Millipedes” (Anonymous, 2003b)
“Introduction to Millipedes” (The Field Museum, 2004)
“Millipedes” (Anonymous, 2003c)
“Polydesmid Millipede Fact File” (Australian Museum, 2003b)
“The Diplopoda (Millipedes)” (Ramel, 2003)
“An Introduction to the Spiders of South East Asia” (Murphy and Murphy, 2000)
“How to know the spiders” (Kaston, 1972)
“Opiliones or Phalangida–what’s in a name” (Curtis, 2003)
“Whip Spiders (Chelicerata: Amblypygi): their biology, morphology and systematics” (Weygoldt, 2000)
“Spider genera of north America” (Roth, 1993)
“The fauna of British India including Ceylon and Burma: Arachnida” (Pocock,
1900)
“The fauna of India. Spider: Araneae” (Tikader, 1982)
“A Field Guide to Insects: America north of Mexico” (Borror and White, 1970)
“An Introduction to the Study of Insects” (Borror et al., 1989)
“The Insects of Australia” (CSIRO Division of Entomology, 1970)
“Antlike Stone Beetle” (Anonymous, 2003a)
“Cerylonidae” (Lawrence, 2003)
“Dermaptera” (Haas, 1996)
“Dermaptera” (Meyer, 2001)
“Featherwing Beetles — Insect: Coleoptera: Ptiliidae” (Dybas, 2000)
“Hymenoptera of the World: An Identification Guide to Families” (Goulet and
Huber, 1993)
“Ptiliidae” (Hall, 1995)
“Rove beetles — Staphylinidae” (Frank and Thomas, 2003)
“A New General Catalogue of the Ants of the World” (Bolton, 1995)
34
Chapter 6. Materials and methods
35
“Australian Ants Online” (Shattuck and Barnett, 2004)
“Identification Guide to the Ant Genera of Khao Yai National Park” (Wivatwithaya and Jaitrong, 2001)
“Identification Guide to the Ant Genera of the World” (Bolton, 1994)
“Identification Guide to Bornean Ants” (Hashimoto, 2003)
“The Ants” (Hölldobler and Wilson, 1990)
“A Field Guide to the Smaller Moths of South-East
Asia” (Robinson et al., 1994)
“A list of Odonata from Thailand (Parts I-XXI)” (Asahina and Pinratana, 1993)
“Atlas of the dragonflies of Thailand: Distribution maps by provinces” (Hämäläinen and Pinratana, 1999)
Classification and nomenclature of higher level taxa follow Borror et al. (1989)
All specimens are deposited in the Forest Insect Group (FIG) Terrestrial Arthropod
collection of the Department of National Parks, Wildlife and Plant Conservation
(DNPW), Thailand.
During sorting specimens were also examined for morphological modification typically associated with hypogean fauna such as reduced or lost eyes, greatly elongated
appendages, reduced pigmentation, etc.
6.4.2
Fish
Field methods
Fish sampling was conducted at only three of the six stations in the end of each cave,
stations 1, 3 and 5 (Figure 6.1). However, at each of these three stations sampling
was conducted in two parts, these parts being upstream and downstream of the central
20 m area that was used for TA sampling. An important consequence of this is that the
first part of the first station was the area immediately outside the cave entrance. In
order to allow more ready comparison with the TA data here we treat each part of the
three fish sampling stations as separate station, thus part 1 of station 1 being regarded
as station 1 while part 2 of station 5 is regarded as station 6. All through the analyses
unless explicitly stated this will be the meaning of ’station’ in relation to fish.
Fish collection was done by two collectors walking and manually searching all water
less than ≈1.5 m deep that fell within the area of each station (20 m before and after
each TA station, Figure 6.1) for a total of 60 minutes, 30 minutes in each section. Fish
encountered were either collected directly with a scoop net or stunned with the aid of
an electro-fishing device and then collected and kept alive in labelled bags and buckets.
Fish identification and morphology
At the completion of the sampling all fish were identified and had their weight and
total length measured. Fish were weighed with either a spring balance (Camry Cap,
2000g) or an electrical balance (Acculab VI-1200). Total length from snout to tail-fin
tip was measured using a standard fish measuring board.
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
After these measurements were completed most fish were returned to the water of
their station. In a few cases where positive identification was not possible in the field
specimens were kept live in aerated labelled plastic bags until they could be fixed and
preserved in formaldehyde for later examination.
Fish morphometric data was mainly used for the calculation of a condition index
(commonly referred to as “condition factor”) based upon the following formula, after
Wootton (1998):
W
k= 3
L
where k, W and L are condition index, fish weight and fish length respectively.
The fish team had many initial sampling problems that badly effected the quantifiability of data from the first sampling trip so the data presented here excludes trip
1.
6.5
Physical and environmental parameters
In addition to the collection of terrestrial arthropods and fish, which are of course
our measures of biodiversity, we were also interested in a wide range of physical and
environmental factors, both because of interest in the factor per se and in their role as
potential co-factors with other variables.
A full description of these variables and their measurement is given here:
Physical characteristics: All caves had the following physical characteristics determined
1. Entrance height: The maximum height of each of the main cave entrances
was either directly measured or estimated to the nearest metre.
2. Entrance width: The maximum width of each of the main cave entrances
was measured.
3. Light penetration: At each station a visual check was made as to whether
any visible light from the entrance reached this point, stations were scored
as either having visible light or not.
4. Station width: At the mid-point of each station the width of the cave was
measured.
5. Station height: At the mid-point of each station the maximum height of the
cave was measured.
6. Maximum flood height: At each station a search was made for deposited
flood debris and other signs of flooding, the maximum height was measured
or estimated in metres.
7. UTM coordinates: The coordinates of each cave’s disturbed entrance were
determined with a Garmin 12 XL GPS.
8. Cave length: Cave lengths were either determined by reference to published
maps or by direct survey using standard cave survey techniques during initial
site survey trips.
36
Chapter 6. Materials and methods
37
9. Cave entrances: The number of cave entrances was determined by reference
to published maps and by direct survey.
10. Catchment area size: Was estimated from 1:50 000 topographical maps. In
the case of KKL the estimate may be subject to considerable error as the
KKL sink and stream does not appear on the map.
Soil samples: During trips two and three small (≈ 30 cm 3 ) surface soil samples were
taken, from each of stations 1, 3 and 5. These samples were analysed by the Soil
Science Department of Kasetsart University (KU) to determine organic matter
content (%). Unfortunately the trip 2 samples were damaged prior to analysis
and these data were lost.
Cave atmosphere: During trips two and three the following characteristics of each
cave’s atmosphere were measured:
1. Air temperature: Wet and dry air temperatures were measured at stations
1, 3 and 5 in each cave, these data being used to determine relative humidity
(%).
2. Draught direction: The direction of any draught was determined by one of
us (DS) as either into or out of the cave.
3. Draught strength: The strength of any detectable draught was determined
by one of us (DS) as being weak, strong or very strong.
Cave Water: During trips two and three the following characteristics of each cave’s
stream water were measured:
1. Temperature: Water temperatures were measured at stations 1, 3 and 5.
2. Flow rate: Water flow rates were measured during trip 3 using a timed float
traveling over a measured distance. During trip 2 an attempt to measure
flow rates with propeller flow meters failed due to low flow rates.
3. Dissolved Oxygen: Dissolved Oxygen (DO) in waters were measured during
trips two and three at stations 1, 3 and 5 using a DO meter (Oxyguard
Gramma/Primatech).
4. Water chemistry: During trip 2, water samples (of 500 ml each) were taken
from stations 1, 3 and 5. These samples were kept dark and relatively cool
prior to analysis by the Watershed Research Laboratory of the DNPW. The
following parameters were determined:
(a)
(b)
(c)
(d)
(e)
(f)
(g)
(h)
9
pH 9
Electroconductivity
Hardness
Na+ concentration
K+ concentration
Ca2+ concentration
Mg2+ concentration
Fe2+ concentration
unfortunately a problem in the laboratory meant that these data were not useful
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
(i) Dissolved Oxygen
Habitat coverage: Estimates were made of the percent coverage of the following
habitat types at each station:
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
Still or slow water
Flowing water
Rapids
Riffles
Boulders
Gravel
Sand
Mud
Bare rock
Flowstone
Concrete
Guano
Flood debris
The estimates were made from sketch maps (see Appendix C) made of each
station during trips 2 and 3. In addition to these directly measured variables
two variables: a) water habitat; b) land habitat were derived. These variables
were calculated by summing habitat types 1-4 to give total water habitat and
obtaining land habitat by substracting the figure from 100%.
No physical or environmental data was collected during trip 1 as it was felt that
during this trip it would be best to concentrate on the main biological data.
6.6
Data management and analysis
Data entry and basic management was done using either Microsoft Excel or Microsoft
Access with further manipulations using the R system (Ihaka and Gentleman, 1996;
R Development Core Team, 2003).
The bulk of data summary, analysis and presentations were also done using R.
Indicator Species Analysis (ISA), by the methods of Dufrene and Legendre (1997),
was done with PC-ORD (McCune and Mefford, 1997).
All data were tested for normality (Gaussian distribution) with the Shapiro-Wilk’s
normality test and heteroscedasticity using the Fligner-Killeen test for homogeneity of
variances prior to analysis. Transformations were applied where appropriate. All logs
used are log-base-2. In cases where data which could not be made Gaussian and/or
homoscedastic by transformation, analysis used Generalized Linear Models which allowed greater flexibility and robustness with data of this type (Venables and Ripley,
1999). In the case of the disturbance scoring data, the non-parametric Wilcoxon paired
test was used because of the ordinate nature of the data.
38
Chapter 6. Materials and methods
39
Sampling effort was examined with plots of species accumulation curves (Colwell
and Coddington, 1994) and singleton and doubleton curves (Scharff et al., 2003), and
calculation of sampling intensity (Coddington et al., 1996) and inventory completion
indices (Sørensen et al., 2002).
We chose to use the first-order jacknife estimator of species richness from among the
many proposed estimators (Colwell and Coddington, 1994). The first-order jackknife
estimator was the only estimator able to provide consistent estimates based upon often
small samples while also being reasonably consistent with other estimators for large
samples. Our observations are similar to those reported by Hellmann and Fowler (1999),
however, they are contrary to those recently reported by Herzog et al. (2002). We are
mindful that whatever estimator used there will be particular problems and are even
more mindful that many “traditional” estimators may give considerable underestimates
(Ugland et al., 2003), though obviously less so than raw species counts.
Canonical Correspondence Analysis (CCA) (Ter Braak, 1986) was used to examine
community composition patterns related to β diversity among caves and disturbance
levels.
Two indices of diversity were chosen, a) Fisher’s α and b) the Shannon-Weiner
index (H0 ). Fisher’s α was adopted following the recommendations of Southwood and
Henderson (2000). The Shannon-Weiner index was chosen despite the many recognised
problems of this index (as summarised in Southwood and Henderson (2000)) because
it is still such a widely used index (especially in Thailand). We hope, perhaps in vain,
that we do not prolong the use of this index by mention of it here.
Because of the problem with Tham Nam 1 only allowing five sampling stations all
analyses presented here are based on stations 1–5. Station 6 data being was dropped
from the analysis.
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
40
Chapter 7
Results
And now for something completely
different.
Monty Python
7.1
Disturbance
Figure 7.1 shows that our a posteriori assessment of relative disturbance levels was supported by quantitative measurement of disturbance levels (one-tailed paired Wilcoxon
test for differences in disturbance scores between disturbed and undisturbed ends: W =
28.0, p = 0.007813).
18
16
10
12
14
Disturbance score
20
22
Disturbed
Undisturbed
Cave Mean
TPN
JKT
KKL
PCH
TNG
NLY
NM1
Figure 7.1: Disturbance scores (mean per station) for each of the seven study caves.
Caves are ordered according to the relative difference in disturbance scores
between ends, this “effects ordering” being recommended for data of this
type (Becker et al., 1996)
Figure 7.1 also shows the range of absolute and relative (difference) disturbance
levels for each of the replicate caves, ranging from Tham Tapan with both the highest
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
overall level of disturbance and the greatest difference between disturbed and undisturbed ends to Tham Nam 1 which had the second lowest level of overall disturbance
and in which there was very little difference in disturbance levels between ends.
7.2
7.2.1
Terrestrial Arthropods
Catch summary and sampling efficiency
Approximately 52 person-hours of cave-time was spent sampling (including travel between stations) in each cave, giving a total of ≈ 364 person-hours of cave-time overall.
A total of 4079 TA specimens were collected representing 519 RTUs from 25 orders
in five classes (Arachnida, Hexapoda (Insecta + Entognatha), Malostraca, Chilopoda,
Diplopoda). Coleoptera proved the most diverse with 98 RTUs, followed by Araneae
(84), Diptera (74), Hymenoptera (56) and Orthoptera (28). The total number of RTUs
collected from each cave was 121 for TPN, JKT = 186, KKL = 94, PCH = 91, TNG
= 130, NLY = 125 and NM1 = 129.
This collecting effort yields a sampling intensity of ≈ 8 while the inventory completion index was 64%. Examining sampling efficiency further with separate species
accumulation curves for each level of disturbance shows that sampling is far from saturated with accumulation curves for the disturbed and undisturbed treatments showing
no indication of reaching an asymptote and the singleton and doubleton curves showing
no sign of intersecting (Figure 7.2).
A total of 456 RTUs had ten or less individuals (≈ 88%) with 250 singletons (≈
48%), 81 doubletons (≈ 16%) and 38 tripletons (≈ 7%).
Another important feature of Figure 7.2, though unrelated to sampling efficiency, is
the clear separation of species accumulation curves for disturbed and undisturbed cave
ends, with the disturbed curve being much higher than the undisturbed one and their
confidence intervals showing no overlap.
42
43
300
200
100
0
Accumulated species
400
Chapter 7. Results
0
100
200
300
400
Sampling effort
Figure 7.2: Randomised TA species accumulation (solid line) with estimated 95%
confidence envelope, singleton (dashed line) and doubleton (dash-dot line)
curves. Blue lines indicate disturbed cave ends, green undisturbed . Curves
based on 100 randomisations, sampling effort is the number of samples
based on a partitioning of disturbance, season, cave and distance.
7.2.2
Abundance
Although strictly the counts of individual animals that we refer to here are indicators
of relative abundance we will use the simpler term abundance throughout this report.
The main factors disturbance, season and cave all showed very significant effects
on TA abundance (Table 7.1). The main effect distance and the interaction terms were
not significant.
The significance of the cave term is uninteresting, in terms of our primary goals
(caves being simply replicates), except in respect to being a clear indication of the value
of the blocked design and the unwanted variation we were able to remove through this
design. As cave effects are largely unimportant Figure 7.3 is the only figure we present
showing cave effects (with a similar figure show for fish), though differences among
cave are significant for all the parameters we tested.
The ends of caves that were disturbed had clearly greater abundances of TA than
undisturbed ends, this being the case regardless of distance and season (Figure 7.4).
Figure 7.4 also shows the season effect with trip 2 TA abundance being lower than trips
1 and 3.
One clearly unusual feature of Figure 7.4 is the pronounced increase in abundance
at station 4 during trip 1 (though note the large confidence interval), with a less
pronounced, but still obvious peak, apparent in trip 2, while no peak is apparent in
the trip 3 data. This peak seems to be “real” and the pattern is seen very clearly in
43
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
three caves (JKT, KKL and TPN). In JKT the pattern is strongest and is seen across
a broad range of taxa while in KKL and TPN it is less marked and results from fewer
taxa with many individuals.
Obviously these rather unusual spikes add significant noise to any trends there
may be for abundance with distance, however, as hinted at by the declining trend
apparent in undisturbed treatment during trip 1 while undisturbed during trip 3 show
an increasing trend there may be no clear trend regardless.
Table 7.1: ANOVA table of effects of disturbance, season and distance on TA abundance
20
15
0
5
10
Abundance
25
30
35
SS
df
F p(>F)
Disturbance
19.11
1 19.28 0.0000
Trip
9.82
2 4.95 0.0081
Distance
1.89
4 0.48 0.7538
Cave
25.74
6 4.33 0.0004
Disturbance × Trip
2.86
2 1.44 0.2394
Disturbance × Distance
5.74
4 1.45 0.2203
Season × Distance
1.80
8 0.23 0.9855
Disturbance × Season × Distance
6.33
8 0.80 0.6044
Residuals
172.50 174
TPN
JKT
KKL
PCH
TNG
NLY
NM1
Caves
Figure 7.3: Overall TA abundance (mean per station ± 95% CI) for each of the seven
study caves
44
45
60
40
0
20
Abundance
80
100
Chapter 7. Results
1
2
3
4
Trip 1
5
1
2
3
4
Trip 2
5
1
2
3
4
Trip 3
5
Figure 7.4: TA abundance (mean per station ± 95% CI) by disturbance level (unfilled:
disturbed ; filled: undisturbed ), season (Trip 1-3) and distance (station
numbers 1-5)
7.2.3
Species richness
Observed species count
Despite the fact that observed species counts are more properly considered as species
density measures rather than measures of richness (Gotelli and Colwell, 2001) we
present observed species counts as it is still common practice to present these data
as they have been traditionally presented.
The significant differences seen with the observed species count are essentially similar to the results seen for TA abundance (Table 7.2). The patterns of differences
in observed species count are also broadly similar to those seen for TA abundance
(Figure 7.5), this, of course reflecting, the lack of sample size dependence problem in
observed species count but also likely reflecting the real situation in the field.
It is interesting to note that the spike seen for station four during trips one and two
is more broadly based for observed species count than abundance, for observed species
count the spike being apparent in five of the caves (JKT, NM1, TPN, NLY and TNG),
but it is not apparent for KKL, unlike abundance.
45
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Table 7.2: ANOVA table of effects of disturbance, season and distance on TA observed
species count
15
10
0
5
Observed species count
20
25
SS
df
F p(>F)
Disturbance
22.23
1 21.12 0.0000
Trip
11.80
2 5.61 0.0044
Distance
4.69
4 1.11 0.3517
Cave
24.68
6 3.91 0.0011
Disturbance × Trip
2.98
2 1.42 0.2456
Disturbance × Distance
3.52
4 0.84 0.5043
Season × Distance
2.58
8 0.31 0.9626
Disturbance × Season × Distance
3.05
8 0.36 0.9391
Residuals
183.10 174
1
2
3
4
Trip 1
5
1
2
3
4
Trip 2
5
1
2
3
4
Trip 3
5
Figure 7.5: TA observed species count (mean per station ± 95% CI) by disturbance
level (unfilled: disturbed ; filled: undisturbed ), season (Trip 1-3) and distance (station numbers 1-5)
Species richness estimates
The species richness estimate (Jacknife 1) again shows significant differences among
distance, season and cave (Table 7.3) as seen for abundance and observed species
counts. The distance effect is also more significant than seen for abundance and observed species counts though not yet significant. Similarly the patterns seen among
disturbance, season and distance (Figure 7.6) are similar to those seen for abundance
and observed species counts.
46
Chapter 7. Results
47
Table 7.3: ANOVA table of effects of disturbance, season and distance on TA estimated species richness
20
0
10
Estimated species
30
SS
df
F p(>F)
Disturbance
29.80
1 30.79 0.0000
Trip
9.26
2 4.78 0.0095
Distance
7.52
4 1.94 0.1057
Cave
20.80
6 3.58 0.0023
Disturbance × Trip
4.26
2 2.20 0.1140
Disturbance × Distance
3.54
4 0.92 0.4563
Season × Distance
2.59
8 0.33 0.9517
Disturbance × Season × Distance
2.55
8 0.33 0.9539
Residuals
168.41 174
1
2
3
4
Trip 1
5
1
2
3
4
Trip 2
5
1
2
3
4
Trip 3
5
Figure 7.6: TA estimated species (mean per station ± 95% CI) by disturbance level
(unfilled: disturbed ; filled: undisturbed ), season (Trip 1-3) and distance
(station numbers 1-5)
7.2.4
Biodiversity indices
Fisher’s α
Fisher’s α estimates proved to be unstable when calculated based on the same model
used for abundance, observed species count and estimated species. As result extra
pooling was required. Pooling was done across distance as distance had proved nonsignificant for other parameters, this means no among distance comparison to be made.
No factors show significant differences in Fisher’s α, though both disturbance and
47
48
The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
season are nearly so (Table 7.4). Though not significant (probably as result of our inability to partition inter-station variation) the patterns seen for disturbance and season
are similar to those seen for other parameters as described previously–disturbed ends
having higher Fisher’s α that undisturbed ends, while trip 2 has a lower Fisher’s α than
other times (Figure 7.7).
Table 7.4: ANOVA table of effects of disturbance and season on TA Fisher’s α
F
3.71
2.77
0.62
0.37
p(>F)
0.0638
0.0787
0.7109
0.6909
2.0
1.5
0.0
0.5
1.0
Fisher’s alpha
2.5
3.0
3.5
SS df
Disturbance
1.15 1
Trip
1.72 2
Cave
1.16 6
Disturbance × Trip 0.23 2
Residuals
9.29 30
Trip 1
Trip 2
Trip 3
Figure 7.7: TA Fisher’s α (mean per station ± 95% CI) by disturbance level (unfilled:
disturbed ; filled: undisturbed ) and season (Trip 1-3)
Shannon Weiner
H0 shows similar results to all the other parameters examined, except Fisher’s α which
is based upon a different model, with disturbance, season and cave effects all significant
(Table 7.5).
48
Chapter 7. Results
49
Table 7.5: ANOVA table of effects of disturbance, season and distance on TA H 0
1.0
0.0
0.5
log2(H’)
1.5
2.0
SS
df
F p(>F)
Disturbance
6.43
1 21.91 0.0000
Trip
1.96
2 3.33 0.0381
Distance
1.06
4 0.90 0.4628
Cave
7.82
6 4.44 0.0003
Disturbance × Trip
0.62
2 1.06 0.3495
Disturbance × Distance
1.81
4 1.54 0.1922
Season × Distance
0.75
8 0.32 0.9577
Disturbance × Season × Distance 1.02
8 0.43 0.8994
Residuals
51.08 174
1
2
3
4
Trip 1
5
1
2
3
4
Trip 2
5
1
2
3
4
Trip 3
5
Figure 7.8: TA H0 (mean per station ± 95% CI) by disturbance level (unfilled: disturbed ; filled: undisturbed ), season (Trip 1-3) and distance (station numbers 1-5)
7.2.5
Relationship between biodiversity parameters and disturbance
scores
In addition to simply testing for a difference between disturbed and undisturbed ends
we also examined whether any more detailed relationships were apparent between all
the biodiversity parameters discussed above and disturbance scores. Plots, correlation
(linear and simple monotonic) and regression were used but no cases were any significant
results found.
49
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
7.2.6
Indicator species analysis
ISA was used to examine which species may be suitable indicators (in the sense of
Dufrene and Legendre (1997)) of disturbance levels and distance.
Indicator species analysis was conducted looking for species that were indicators
of disturbance or distance (neither season or cave were examined as the former is by
definition likely to be only ephemeral (and for which we have no seasonal level of
replication) while the later simply shows the large among cave differences that are
expected).
Only one RTU was found to be a significant (at α< 0.05) indicator of disturbance,
while two other RTUs were marginally significant (at 0.05 <α< 0.1). These RTUs were
indicators of the disturbed ends of caves.
Table 7.6: Summary of TA ISA scores and their significance for disturbance
Class
Order
Family
Taxa code
Arachnida
Diplopoda
Hexapoda
Araneae
Julida
Coleoptera
Ochyroceratidae
—
Staphylinidae
Aran052
Diplopoda sp15
Col0093
p
0.063
0.085
0.024
disturbed
indicator value
57
57
72
undisturbed
indicator value
0
0
2
Nine RTUs were found to be significant as indicators of distance, with a further two
RTUs marginally significant”. Two RTUs were found to be indicators of deeper stations
(stations 4 and 5) while the remainder were indicators of cave entrances, mostly Station
1, however, one predatory reduviid was associated with station 2.
Table 7.7: Summary of TA ISA scores and their significance for distance
Class
Order
Family
Taxa code
Arachnida
Arachnida
Diplopoda
Hexapoda
Hexapoda
Hexapoda
Hexapoda
Hexapoda
Hexapoda
Hexapoda
Malostraca
Araneae
Araneae
Glomerida
Hemiptera
Hymenoptera
Hymenoptera
Hymenoptera
Coleoptera
Coleoptera
Coleoptera
Isopoda
Nesticidae
Salticidae
—
Reduviidae
Formicidae
Formicidae
Formicidae
Staphylinidae
Staphylinidae
—
—
Aran028
Aran025
Diplopoda sp7
Emesinae001
For025
For022
For021
Col0093
Col0032
Col0004
Isopoda sp1
50
p
0.036
0.001
0.097
0.005
0.005
0.018
0.001
0.047
0.078
0.029
0.024
Station Indicator Value
1
2 3 4
5
36 2 0 0
0
86 0 0 0
0
0
0 6 0 24
0 49 0 0
2
57 0 0 0
0
43 4 0 0
0
86 0 0 0
0
33 6 0 0
0
35 1 0 1
0
1
4 0 35 2
43 0 0 0
0
Chapter 7. Results
7.2.7
51
Community composition
Community ordination
The clearest pattern evident in the CCA ordination plot is the similarity of disturbed
and undisturbed ends within the same cave (Figure 7.9). Of course, this is hardly
surprising as we have seen in earlier analyses that cave factor is also important in other
aspects of our data. Aside from the ends of caves there also seems to be an indistinct
regional clumping effect. All the Phangnga cave ends (TPN, PCH, TNG and NM1),
except the disturbed end of TPN which is quite far from most other cave ends, form
a tight cluster near the origin. The Satun caves (JKT and KKL) also showing some
similarities.
KKL
1
2
KKL
0
NM1
PCH
PCH TNG
NM1 TNG
TPN
JKT
NLY
NLY
−2
−1
CA2
JKT
TPN
−4
−2
0
2
4
CA1
Figure 7.9: TA CCA ordination plot showing the 14 study cave ends. Blue: disturbed ;
Red: undisturbed
Taxonomic composition at the Order level
In addition to ordination we examined community composition at the Order level using Mosaic plots (Friendly, 1994) to test the independence of community composition
and disturbance levels. We examined the ten most abundant Orders. The most abundant Order was Araneae, comprising ≈ 41% of the specimens, with other major orders
being Orthoptera (≈ 15%), Coleoptera (14%) and Hymenoptera (≈ 10%), these four
orders representing around 80% of specimens (Figure 7.10). Of these major orders
Araneae and Orthoptera (being dominated by Rhaphidophoridae) were both overrepresented (in relation to expectations) in undisturbed cave ends while Coleoptera
was over-represented in disturbed ends, and Hymenoptera showed no significant bias
(Figure 7.10).
51
52
0:2
2:4
>4
Orthoptera
Isopoda
Opiliones
Hymenoptera
Dermaptera
Diplopoda
Diptera
Hemiptera
Coleoptera
Araneae
The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Standardized
Residuals:
<−4
−4:−2
−2:0
Disturbed
Undisturbed
Figure 7.10: Mosaic plot of order level taxonomic composition differences between
disturbance levels
7.2.8
Morphological modifications for hypogean life
Many RTU showed morphological modifications typical of troglobytes such as absent
or reduced eyes (Diplopoda and Araneae), elongated appendages (Amblypygida, Orthoptera), etc. However, none showed sufficient modification for us to regard them as
troglobytes, most likely being troglophiles with morphology pre-adapted to cave life.
7.3
7.3.1
Fish
Catch summary and sampling efficiency
A total of 1681 specimens were collected representing 57 species from 40 genera in
17 families (full taxonomic details in Appendix B.1). These figures giving a sampling
intensity of ≈ 29 and inventory completion index of 78%.
We have found a species which expected to be a new species and one species is a
new record for Thailand.
Examining sampling efficiency in detail for each level of disturbance shows that
both disturbed and undisturbed levels had reasonably saturated sampling as indicated
by the approaching of an asymptote by the species accumulation curves and the fact
52
Chapter 7. Results
53
30
20
0
10
Accumulated species
40
50
that the curves for singletons and doubletons for both levels of disturbance have crossed
(Figure 7.11). Figure 7.11 also shows a clear separation of species accumulation curves
for disturbed and undisturbed cave ends, with the disturbed curve being much higher
than the undisturbed one and their confidence intervals showing no overlap.
Pooling the entire sample (both levels of disturbance) shows 11 singletons (≈ 19%),
12 doubletons (≈ 21%) and 2 tripletons (≈ 4%). A total of 40 species had ten or less
individuals (≈ 70%).
0
20
40
60
80
Sampling effort
Figure 7.11: Randomised fish species accumulation (solid line) with estimated 95%
confidence envelope, singleton (dashed line) and doubleton (dash-dot
line) curves. Blue lines indicate disturbed cave ends, green undisturbed .
Curves are based on 100 randomisations, sampling effort is the number
of samples based on a partitioning of disturbance, season, cave and
distance.
7.3.2
Abundance
The main effects disturbance, distance and cave all showed significant differences, while
there was no significant difference between season (Table 7.8). None of the interaction
terms were significant.
As with TA cave term is interesting only in terms of indicating the effectiveness of
the blocked design. Examination of Figure 7.12 makes it also seem likely that a fair
deal of the inter-cave variation results from catchability differences among cave streams
as it is noteworthy that the two largest streams (JKT and NLY, more like small rivers)
had the lowest abundance and while this may be a reflection of the actual abundance
the size of these streams also made catching fish difficult and so this may reflect this
53
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
fact. Figure 7.12 is the only figure we present showing cave effects for fish though
differences among cave are significant in most cases.
With respect to the effects of disturbance and distance, Figure 7.13 shows clearly
that disturbed cave ends had greater abundance than undisturbed ends and that abundance was much greater at the entrance or short distances from it, dropping off sharply
with distance from the entrance.
Table 7.8: ANOVA table of effects of disturbance, season and distance on fish abundance
3.0
2.5
2.0
1.5
1.0
0.0
0.5
Fish abundance (log2(abundance))
3.5
SS df
F p(>F)
Disturbance
7.05 1 4.24 0.0424
Season
0.75 1 0.45 0.5041
Distance
109.29 4 16.45 0.0000
Cave
28.99 6 2.91 0.0125
Disturbance × Season
0.51 1 0.31 0.5822
Disturbance × Distance
4.78 4 0.72 0.5810
Season × Distance
5.51 4 0.83 0.5103
Disturbance × Season × Distance
2.19 4 0.33 0.8578
Residuals
141.17 85
TPN
JKT
KKL
PCH
TNG
NLY
NM1
Cave
Figure 7.12: Overall fish abundance (mean per station ± 95% CI) for each of the
seven study caves
54
55
4
3
2
0
1
Fish abundance (log2(abundance))
5
6
Chapter 7. Results
1
2
3
Trip 2
4
5
1
2
3
Trip 3
4
5
Figure 7.13: Fish abundance (mean per station ± 95% CI) by disturbance level (unfilled: disturbed ; filled: undisturbed ), season (Trip 2-3) and distance
(station numbers 1-5)
When abundance is considered with respect to inflow versus outflow we find that
inflow/outflow had no effect on fish abundance, though distance and cave effects were
again significant (Table 7.9 and Figure 7.14).
Inflow/outflow was also found to be not significant for any of the other parameters
we examined and as a result it was not necessary to adopt a block with regard to this
effect and we examine this factor no further in these analyses.
Table 7.9: ANOVA table of effects of inflow-outflow, season and distance on fish abundance
SS df
F p(>F)
Flow
2.48 1 1.47 0.2282
Season
0.64 1 0.38 0.5386
Distance
103.47 4 15.34 0.0000
Cave
24.26 6 2.40 0.0345
Flow × Season
0.77 1 0.45 0.5020
Flow × Distance
4.36 4 0.65 0.6305
Season × Distance
6.25 4 0.93 0.4524
Flow × Season × Distance
4.87 4 0.72 0.5790
Residuals
143.31 85
55
The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
4
3
2
1
0
Fish abundance (log2(abundance))
5
56
1
2
3
Trip 2
4
5
1
2
3
Trip 3
4
5
Figure 7.14: Fish abundance (mean per station ± 95% CI) by inflow-outflow (unfilled:
inflow; filled: outflow), season (Trip 2-3) and distance (station numbers
1-5)
7.3.3
Species richness
Observed species counts
As was seen with abundance the significant main effects were disturbance, distance
and cave. The season main effect and the interaction terms were not significant (Table 7.10).
Not surprisingly the effects of disturbance and distance on observed species counts
were similar to those seen for abundance with disturbed cave ends having higher observed species counts than undisturbed ends, while stations close to the entrance, especially the first station, immediately outside the cave entrance, had higher observed
species counts than stations more distant from the entrance (Figure 7.15).
56
Chapter 7. Results
57
Table 7.10: ANOVA table of effects of disturbance, season and distance on fish observed species count
5
4
3
0
1
2
Observed species count
6
7
SS
df
F p(>F)
Disturbance
50.40
1 11.02 0.0012
Season
4.83
1 1.06 0.3064
Station
149.39
4 8.17 0.0000
Cave
62.89
6 2.29 0.0399
Disturbance × Season
0.00
1 0.00 1.0000
Disturbance × Station
5.39
4 0.29 0.8811
Season × Station
18.10
4 0.99 0.4164
Disturbance × Season × Station
2.50
4 0.14 0.9684
Residuals
521.40 114
1
2
3
Trip 2
4
5
1
2
3
Trip 3
4
5
Figure 7.15: Fish observed species count (mean per station ± 95% CI) by disturbance
level (unfilled: disturbed ; filled: undisturbed ), season (Trip 2-3) and
distance (station numbers 1-5)
Estimated species richness
In order to get “reliable” species estimates it is necessary to have reasonable numbers
of individuals in each sample. Unfortunately this meant that rather than being able
to examine all the main effects examined previously viz. disturbance, season, distance
and cave it was necessary to drop one factor to allow pooling in order to have sufficient
data to make reliable estimates. As season had shown no significant effects for either
abundance or species density it was dropped.
The main effects disturbance and distance showed significant effects, while cave was
57
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
not significant (Table 7.11).
As seen with abundance and observed species counts disturbed ends had greater
estimated species richness than undisturbed ends and stations at or near the entrance
had higher species richness estimates than more distant stations, although both station
three of the undisturbed caves had much higher levels of species richness than might be
expected given their locations (Figure 7.16).
Table 7.11: ANOVA table of effects of disturbance, season and distance on fish estimated species richness
F
9.09
6.70
1.12
0.08
p(>F)
0.0039
0.0002
0.3639
0.9877
8
6
0
2
4
Estimated species
10
12
14
SS df
Disturbance
111.89 1
Station
330.09 4
Cave
82.65 6
Disturbance × Station
4.02 4
Residuals
664.92 54
1
2
3
4
5
Station
Figure 7.16: Fish estimated species (mean per station ± 95% CI) by disturbance level
(unfilled: disturbed ; filled: undisturbed ) and distance (station numbers
1-5)
58
Chapter 7. Results
7.3.4
59
Biodiversity indices
Fisher’s α
Like species richness estimators in order to obtain numerically stable estimates of
Fisher’s α it was necessary to clump over the season effect. In addition several estimates of Fisher’s α were still clearly unstable and so prior to analysis all Fisher’s α
estimates exceeding 100 were arbitrarily assigned as missing data (NA), this effectively
meant that values exceeding 8195.26 were excluded (8 cases in all), this being the next
value over 100.
The only significant main effect was disturbance, cave showing no significant effect
(Table 7.12). As seen with the other parameters again it was the disturbed cave ends
that had higher Fisher’s α values than undisturbed ends.
Table 7.12: ANOVA table of effects of disturbance, season and distance on fish
Fisher’s α
F
5.77
1.42
1.65
1.28
p(>F)
0.0212
0.2439
0.1604
0.2957
6
4
0
2
Fisher’s alpha
8
10
SS df
Disturbance
57.03 1
Station
56.35 4
Cave
97.68 6
Disturbance × Station 50.48 4
Residuals
385.59 39
1
2
3
4
5
Station
Figure 7.17: Fish Fisher’s α (mean per station ± 95% CI) by disturbance level (unfilled: disturbed ; filled: undisturbed ) and distance (station numbers 1-5)
59
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Shannon Weiner
H0 , unlike Fisher’s α, more closely reflects the patterns seen in abundance, observed
species count and estimated richness, with again disturbance and distance showing
significant effects, though again cave was not significant (Table 7.13).
Again also, we see the familiar pattern of disturbed ends of caves having higher H 0
than undisturbed ends while outer stations, particularly the entrance, had higher H 0
than inner stations (Figure 7.18).
Table 7.13: ANOVA table of effects of disturbance, season and distance on fish H 0
0.0
0.5
H’
1.0
1.5
SS
df
F p(>F)
Disturbance
3.28
1 10.01 0.0020
Season
0.13
1 0.40 0.5281
Station
7.17
4 5.47 0.0005
Cave
4.01
6 2.04 0.0660
Disturbance × Season
0.03
1 0.10 0.7557
Disturbance × Station
0.29
4 0.22 0.9244
Season × Station
1.28
4 0.98 0.4237
Disturbance × Season × Station 0.34
4 0.26 0.9042
Residuals
37.32 114
1
2
3
Trip 2
4
5
1
2
3
Trip 3
4
5
Figure 7.18: Fish H0 (mean per station ± 95% CI) by disturbance level (unfilled:
disturbed ; filled: undisturbed ), season (Trip 2-3) and distance (station
numbers 1-5)
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Chapter 7. Results
7.3.5
61
Relationship between biodiversity parameters and disturbance
scores
As with the TA data we looked for relationships between disturbance scores and the
biodiversity parameters discussed above. No significant results were found.
7.3.6
Indicator species analysis (ISA)
Only one species, Lepidocephalichthys birmanicus (Rendhal), was found to be a significant (at α< 0.05) indicator of disturbance, with another, Rasbora paviei (Tirant), being
marginally significant (at 0.05 <α< 0.1) (Table 7.14) . Both species were indicators of
disturbed cave ends.
Table 7.14: Summary of Fish ISA scores and their significance for disturbance
Species
p
Lepidocephalichthys birmanicus
Rasbora paviei
0.018
0.083
disturbed
undisturbed
indicator value indicator value
36
1
42
8
Seven species were found to be significant (at α< 0.05) indicators of distance (Table 7.15). All seven species were indicators of the cave entrances. All the species feed
on zooplankton and terrestrial insects at the surface of water except S. magnifluvis
feeds mainly on benthic organism.
Table 7.15: Summary of Fish ISA scores and their significance for distance
Species
Brachydanio kerri
Danio aequipinnata
Danio regina
Rasbora paviei
Schistura magnifluvis
Systomus binotatus
Systomus lateristriga
7.3.7
p
0.020
0.006
0.002
0.001
0.036
0.002
0.001
Station
1 2
24 6
22 3
27 5
29 1
5 15
35 5
27 1
Indicator Value
3 4 5
1 1 0
0 0 1
2 1 3
1 1 0
0 0 1
6 1 3
0 0 0
Community composition
The pattern most evident in the CCA of the fish communities is the similarity of
disturbed and undisturbed ends of caves (Figure 7.19). This pattern is not surprising as
we have already seen that the cave factor is also important in other aspects of our data.
The next most obvious pattern is a regional clumping with the Satun caves (KKL and
JKT) forming a cluster, while the spatially isolated NLY shows a clearly distinct fish
fauna. The Phangnga caves are a little unusual as they fall into two groups, a small
group of three cave ends, consisting of disturbed TPN and PCH and undisturbed NM1,
while the remainder form another group. The distinct fauna between ends for TPN and
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
PCH is not surprising as these caves had quite large differences in disturbance between
ends but NM1 ends were very similar so why the fish fauna should be so distinct is not
clear.
2
JKT
KKL JKT
1
KKL
CA2
TPN
0
PCH
NM1
NLY
−1
TNG
NLY
TNG
NM1
TPN
PCH
0
1
2
3
CA1
Figure 7.19: Fish CCA ordination plot showing the 14 study cave ends. Blue: disturbed ; Red: undisturbed
7.3.8
Condition index changes
We were interested in whether fish showed any change in condition, perhaps as a result
of our energy theory, between season. We examined the k and body length of all
those species with more than 10 individuals found in both disturbance levels, this was
coincidentally a total of ten species.
We used ANOVAs of k and body length to examine disturbance, season and their
interaction (while blocking for caves and merging distance). We included body length as
a check for phenological or other season changes in body form that could be confounded
with condition.
Of the ten species several showed changes in k associated with either season, disturbance or the interaction, but only one species, Systomus binotatus (Valenciennes),
showed significant changes in k without any changes in length (Table 7.16). Figure 7.20
62
Chapter 7. Results
63
shows the season and disturbance effects for S. binotatus. S. binotatus in both disturbed
and undisturbed cave ends shows a decline with season, being most obvious in the undisturbed ends where initially the condition index of S. binotatus is undisturbed was higher
than disturbed , but by the final sampling it has declined below that of disturbed . This
general pattern was also evident for most of the other species examined, though as the
results were either no significant or may be a result of confounding factors they are not
show here.
Table 7.16: Summary of ANOVA (p-values) for the condition index k and body length
of ten of the more common fish species
Species
Trip
0.5843
0.1994
0.9291
0.2702
0.4971
0.0018
0.4678
0.1462
0.0003
0.0329
0.014
0.012
0.008
0.010
Condition index
0.016
0.018
Brachydanio kerri
Channa gachua
Danio aequipinnata
Danio regina
Mystacoleucus marginatus
Rasbora paviei
Schistura magnifluvis
Silurichthys schneideri
Systomus binotatus
Systomus lateristriga
Condition Index
Length (cm)
Disturbance Season × TT
Trip
TT
Trip:TT
0.3721
0.0206
0.0250 0.2494
0.0873
0.0743
0.1130
0.2515
0.1099
0.8909
0.3816
0.0373
0.0001 0.5136 0.0044
0.3014
0.9984
0.0521
0.5359
0.1533
0.2462
0.0234
0.6888 0.0046 0.0265
0.5672
0.1052
0.0001 0.9630 0.0001
0.9689
0.2395
0.6168
0.2899
0.7648
0.5429
0.6902
0.0202 0.3174
0.3488
0.2266
0.0358
0.0195 0.0020
0.3001
0.0422
0.0040
0.2694
0.2621
0.5703
1
2
3
Season
Figure 7.20: Changes in Systomus binotatus condition index (mean ± 95% CI) with
season. Unfilled: disturbed ; Filled: undisturbed
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7.3.9
Morphological modifications for hypogean life
None of the specimens collected showed any morphological modification to hypogean
life.
7.4
Physical and environmental data
Physical and environmental data were collected with the purpose of both providing a
simple description of the caves and more importantly as a set of co-factors to be tested
as to whether they were either possibly affected by human disturbance and/or were
confounded with it.
Table 7.17 shows summary of the most important physical and environmental parameters for each of the caves.
Two approaches were used to examine all the physical and environmental variables
collected. The first approach was a set of simple ANOVAs along the same lines as
for the biological data presented above. Of the thirty eight variables examined none
were found to show any significant difference between disturbed and undisturbed levels
of disturbance (thus alleviating any comparison-wise error rate concerns). In addition to this simple approach the tree-based partitioning approach known as recursive
partitioning (Therneau and Atkinson, 1997; Venables and Ripley, 1999) was used to
determine if any of the physical or environmental variables were “predictors” of disturbance— the finding of a “predictor” being important as it may be a confounding factor
for disturbance. No variable was found to be a significant predictor of disturbance.
Table 7.17: Environmental and physical characteristics of the seven study caves
Cave
TPN
JKT
KKL
PCH
TNG
NLY
NM1
a
b
Length
(m)a
1 200
719
757
1 150
960
1 095
640
Altitude
(MASL)
20
15
35
10
80
80
90
Catchment
area (km2 )
1.1
112
≈4
1.1
1.3
110
5.3
Air temp.
(◦ C)b
24.5 - 25.3
25.5 - 26.8
25.0 - 26.8
25.0 - 25.5
24.5 - 26.0
26.4 - 27.0
25.3 - 26.3
Water temp.
(◦ C)b
24.5 - 26.0
25.5 - 26.2
25.0 - 26.2
25.0 - 26.0
24.5 - 25.9
27.5 - 29.1
25.5 - 25.7
Relative
humidity (%)b
97 - 99
83 - 96
96 - 99
96 - 100
96 - 99
97 - 100
93 - 98
Flow rate
(m3 s-1 )
0.0012
1.41
0.03
NA
0.00085
3.605
0.081
the total, combined length of all surveyed passages (taken from published sources)
the range of trip averages
64
Entrances
3
3
4
4
3
3
3
Chapter 8
General discussion
The whole is more than the sum of
the parts.
Metaphysica
Aristotle
8.1
8.1.1
The parts
Sampling effectiveness
Unsurprisingly TA and fish sampling showed different levels of efficiency, fish sampling
being quite complete while TA sampling being far less so.
Judging by the continuing steep ascent of the TA species accumulation curves, the
ascent of the singleton and doubleton curves without any crossover, as well as the
high rate of singletons, TA sampling is far from complete. The sampling intensity and
inventory completion index support this conclusion. Of course, this situation is rather
typical for tropical TAs, indeed TAs in many habitats. Stork (1995) stated bluntly
that “accumulation curves for insects from forests never reach an asymptote”. Scharff
et al. (2003) consider a minimum sampling intensity of 30 to be required for reasonably
reliable species richness estimate, though neither they, nor many others have achieved
this. Scharff et al. (2003) themselves only managed a sampling intensity of 12 (after
correction). It is also worth considering that these sampling efficiencies are measured
at the pooled disturbance level, at lower levels sampling efficiency is likely to be less.
The lack of complete sampling means that this study joins the almost two dozen
“state of the art” arthropod surveys reviewed in Scharff et al. (2003) suffering from
the problem of undersampling bias. This certainly means that care should be taken
interpreting richness and biodiversity measures as absolute estimates. However, the fact
that this study has true replication means that we are able to provide more reliable
estimates of parameters and more importantly of their confidence intervals. We are
quite confident that although the sampling is incomplete our conclusions are entirely
valid — and certainly more valid than other undersampled and unreplicated TA studies.
The desire for complete sampling to ensure reliable species richness estimates (the
estimation of which is strongly based upon reducing the number of singletons) is likely
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to be polemic with regard to sampling cave faunas as these have very low abundance
rates and for which there are considerable dangers of over-collecting (Culver, 1982).
With respect to fish all the measures discussed above show that fish sampling is
quite complete, most noticeably the cross-over of singleton and doubleton curves and
the sampling intensity of 29 — almost reaching the “magic” 30 of Scharff et al. (2003).
8.1.2
Catch summary
The total number of TA RTUs found by this study was 519. The number of TA
RTUs per cave ranged from 91 to 186. Whilst recognising that comparisons of total
species numbers between caves and between studies is not particularly useful due to
different objectives, sampling techniques, cave ecosystems, taxonomic capability, etc.,
for completeness the following studies can be cited. McClure (1965) identified 158
invertebrate species and found more that were left unidentified in Batu Caves, Malaysia.
This number included several aquatic species.
Suhardjono (2003) caught many more species during the course of their study of
cave biodiversity in Sulawesi, Indonesia, than were encountered during this study. This
is most likely due to the different collecting techniques and experimental design adopted
by each study. It is probably not due to Indonesian caves having a naturally greater
biodiversity than caves in Thailand. Evidence for this is given by a comparison between
Gua Sulakkan Kallang, Indonesia, and Tham Chiang Dao, northern Thailand, where
the former contains 93 species and the latter 92 (Deharveng and Bedos, 2000). This
same reference also gives a total species number for two other Indonesian caves: Ngalau
Surat = 74 species and Batu Labang = 72.
8.1.3
Disturbance effects
Both TAs and fish show clear effects from disturbance. TAs and fish both show higher
levels of abundance, richness (observed species counts and estimated species) and α
biodiversity measures (Fisher’s α and H0 ) in disturbed cave ends. These effects are
apparent regardless of season or distance.
Are these results unusual for TAs? The effects of disturbance on TA biodiversity,
and biodiversity generally, are far from clear. With respect to TA some studies have
reported reduced biodiversity (Greenberg and Forrest, 2003; Haskell, 2000; Hill et al.,
1995; Holloway et al., 1992) while others have found the opposite or no effect (Davies
et al., 1999; Eggleton et al., 1996, 1995, 1997; Intachat et al., 1997; Spitzer et al., 1997;
Van Horne and Bader, 1990). Even others have claimed differences, often in the nature
of reduced biodiversity associated with disturbance, while their own results do not seem
to support their conclusions. Good examples of this latter category are Lawton et al.
(1998) and Goehring et al. (2002). Lawton et al. (1998) state that “species richness
declines with increasing habitat modification”, however, examination of their Figure
1 shows only one taxa that seems to show any clear decline. Furthermore this paper
fails to test the significance of any of the supposed changes, perhaps because they
realised too late, that their “replicates” are not valid replicates, but rather are “pseudoreplicates” sensu Hurlbert (1984). Unfortunately methodological failings like this
66
Chapter 8. General discussion
67
seem to be common in studies looking at the effects of disturbance on TA biodiversity,
pseudoreplication is common, no attempt at replication probably more common (i.e.
Jones et al. (2002)), and occasionally, if an experimental approach is taken, failure to
properly disperse treatments (i.e. Davies et al. (1999)) — it seems many TA ecologists
are still repeating many of the methodological problem highlighted by Hurlbert (1984)
twenty years ago.
Aside from methodological failings, there are many reasons why the effects on TA
biodiversity of disturbance have proved to be confused. We mention briefly some the
more obvious ones here:
Different taxa show different responses: Not surprisingly different taxa respond
to disturbance in different ways, some showing increased biodiversity in the presence of disturbance, while others show reductions. The only surprising issue is
that many biologists seem to overlook this fact.
Responses need not be monotonic: The observations that led to the development
of the Intermediate Disturbance Hypothesis (IDH) were that at low levels of habitat disturbance increasing disturbance was associated with increased biodiversity,
but at higher levels of disturbance extra disturbance lead to decreased biodiversity, so producing a humped distribution with maxima at “intermediate” levels
of disturbance (Connell, 1978). There is also evidence of a humped distribution
of species richness along productivity gradients (which could well correlate with
disturbance) (Waide et al., 2000). Obviously sampling over disturbance levels in
the lower part of the curve shows increasing biodiversity while sampling at higher
levels of disturbance leads to the opposite conclusion.
Spatial scale is important: In addition to the possibility of the scale over which
disturbance varies being potentially important the spatial scale of the disturbance,
or the spatial scale of sampling, or the spatial scale over which we interpret our
conclusions can be important. Despite the widely used terms α-biodiversity, βbiodiversity, γ-biodiversity, etc surprisingly few biologists understand that high
α-biodiversity does not mean that β-biodiversity must be high 1 . Furthermore
the lines (spatial scale) between α and β biodiversity are not always clear adding
to the confusion.
Not letting the data speak for themselves: As mentioned above, some authors,
e.g. Goehring et al. (2002); Lawton et al. (1998) add to the confusion by making
claims that their data do not support, or indeed may refute.
Is the situation with respect to data for caves clearer? As there is remarkably
little quantitative data - the situation is certainly simpler. The “common wisdom”
based upon casual observations and isolated case-studies is that disturbance leads to
catastrophic reductions in biodiversity levels in caves. Often, disturbance does result
in declines and extinctions of cave fauna and many examples of such situations have
been described, e.g. Chapman (1993); Hamilton-Smith and Eberhard (2000) and Elliot (2000). However, even here there are exceptions, Spate and Hamilton-Smith (1991)
1
and this situation seems likely to worsen as lecture notes and other materials aimed at students
are equally confused, a good example being Williams et al. (1988) where they are clear on the nonadditivity of scales of biodiversity in some places but not in others!
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
cited a case in Dog Leg Cave where erosion, modified stream flow and visitors “wallowing” in pools has not had a disastrous effect on the relic Gondwanaland invertebrates
found in the cave.
We can cite two quantitative studies which present some data on the disturbance
effects on cave TAs. Ferreira and Horta (2001) surveyed seven caves in the Peruaçu
River valley of Brazil. One cave (Bonita cave) was regarded as a cave heavily disturbed
by tourists, the species richness for Bonita cave was 14 (shared by one other cave), while
the range for the five dry caves in this study was 14–16. In our sister project’s study of
the Maros karst, Sulawesi, Indonesia, they found that Gua Mimpi, which they regarded
as being the most disturbed had the highest diversity (although a less disturbed cave
Gua Togendra had the highest richness) (Suhardjono, 2003). In both these cases there
is no compelling data to suggest reduced biodiversity in the presence of disturbance.
The findings of Suhardjono (2003) being similar to our own.
The fish species studied by this project show same tendency as TAs, that is they
prefer to live in disturbed areas rather than undisturbed and that disturbance has led
to increased biodiversity.
As far as we are aware, there have been no previous reports of studies of human
impacts on trogloxene cavefish. In general, ichthyologists who study cave fish tend
to concentrate on troglobitic animals and their adaptations to cave life, population
densities and reproductive strategies and not the impacts caused by people. For example Brown et al. (1998) extensively surveyed the troglobitic fish populations and
environmental quality in Cave Springs Cave, Arkansas, USA. However, it is known
that cave fish (both trogloxenic and troglobitic) are very susceptible to environmental
changes and lights (Helfman et al., 1997; Parzefall, 1993). Some possible evidence for
environmental degradation in the Phangnga karst has already been found in the distribution of cave dwelling, Balitorid and Cobitid loaches which are very sensitive to
impact (Borowsky, 1998). The more heavily visited caves located near urban centres
and having catchment areas largely turned over to agriculture lacked these fish. Less
impacted caves contained flourishing colonies.
How can disturbance result in higher abundance, richness and diversity in caves
Elliot (2000); Hamilton-Smith and Eberhard (2000) give several sources of threats
to cave fauna. These include: hydrological changes, quarrying, land development,
agriculture, chemical pollution, introduction of exotic species and visitors to caves.
Most, if not all of these sources of impact would be expected to reduce biodiversity of
caves and not increase it. The results of this study therefore contradict the traditional
beliefs that disturbance leads to extinction and decline.
Caves, even high-energy caves, are characterised by low energy availability when
compared to “outside” habitats (Spate and Hamilton-Smith, 1991). The result of this
is what Spate and Hamilton-Smith (1991) referred to as the “Paradox of Enrichment”
whereby the enrichment of the energy supply to a cave can have considerable effects on
cave fauna communities. Elliot (2000) states that this effect is a little studied, subtle
process. It seems likely that higher abundances of TAs and fish in the study is the result
of the introduction of increased energy entering into disturbed cave ends. In some cases
this may be subtle such as the introduction of organic matter in the form of lint or
mud on shoes and clothes (however a test of soil organic matter did not detect any
68
Chapter 8. General discussion
69
significant difference between disturbed and undisturbed ends) or Photosynthetically
Active Radiation (PAR) from electric lights. While in other cases it may be less subtle
such as feeding of fish in temple grounds2 or insects being attracted into caves by
electric lights which are typically left burning continually. Other unnatural sources
of energy include dropped litter and food stuff and use of biodegradable materials in
tourist infrastructure. Of course, higher energy availability may also mean increased
niche availability and associated higher richness and diversity.
In addition to being fed, fish associated with temple caves (always at disturbed
ends) are likely to be less subject to fishing by locals as being on temple grounds
usually means animals will not be hunted. In Tham Tapan for example there was a
“No Fishing” sign. We, indeed were concerned about collecting fish and TA on temple
grounds but discussions with temple monks (at Tham Tapan, Tham Phung Chang and
Tham Thong) seemed to indicate that it was “fine [not sinful] for scientific purposes”.
A further possible mechanism leading to greater richness and diversity, and perhaps abundance, in disturbed cave ends is the accidental introduction of epigean fauna.
Again this may be subtle, in the form of insects traveling into caves on the clothes of visitors, or more direct such as the introduction of fish that are released into caves as part
of Buddhist merit making practices3 . A good example of the latter being the collection
of exotic fish species in the disturbed end of Tham Phung Chang. Reeves (2002) also
discusses the danger of the introduction of exotic species, particularly invertebrates,
into caves in the US.
8.1.4
Season effects
The response of TAs and fish to season was different. TAs showed an effect due to
season while fish did not.
TAs showed markedly lower abundance, richness and biodiversity during the middry season (trip 2). This was certainly not in accord with our initial hypothesis that
seasonal changes in energy availability in the cave would be important. A mid-dry
season minima for energy levels does not seem very likely and so our data seem to be
inconsistent with our ideas about changes. It is most unfortunate that our mid-dry
season organic matter samples were destroyed as these would have provided vital data
on this issue.
The realisation that many of the TA are accidental epigeans (or perhaps trogloxenes)
has led us to consider a new hypothesis. We postulate that the mid-dry season minima
is merely a reflection of the conditions outside the caves. During the middle of the
dry season epigean TA abundance and activity is at a seasonal low. We believe the
relatively low TA abundance and activity outside of the caves results in less accidental
entry of TAs and so the changes seen in the cave fauna merely reflect seasonal changes
outside of caves. If this theory is correct, and it seems to be consistent with our data
but needs proper testing, it casts doubt on the widespread belief that caves faunas are
relatively unaffected by seasonal changes. Of course, clearly with seasonally flooded
2
at the ARCBC Regional Research Grant Conference: Building bridges between ASEAN and EU
researches[sic] (Bangkok, December 1-4 2003) Edmund Gittenberger neatly summed up the situation
with his comment: “ah yes the fish they like that”
3
a practice recognised as particularly dangerous in terms of the introduction of invasive alien
species and one which the Thai government has tried to control
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
caves this will be less so than dry caves but this season effect can obviously affect dry
caves and may be even more important for them. Shaw and Davis (2000) also show a
seasonal (summer versus winter) effects on cave invertebrates from caves on Vancouver
Island, Canada.
Fish showed no response to season, though season will affect catching ability. High
water levels and turbid water during the rainy season will reduce the visibility of fish
and directly affect the catching result, as occurred during trip 3 at NLY cave. Aside
from this, strong currents will sweep fish from cave inflows to outflows. Almost all
tropical fish reproduce in the rainy season, as evidenced by nuptial tubercles (strong,
rigid organs in the cheek and snout of mature fish) and full egg sacs in female specimens
from NLY samples.
Although fish biodiversity parameters showed no effect due to season one species
of fish showed changes in condition with season. S. binotatus showing a decline in
condition from the start of the dry season to the end of the dry season. Interestingly
the decline was sharpest in undisturbed cave ends and much less in disturbed cave ends.
This difference may be an indication that the energy enrichment that we believe occurs
in disturbed cave ends helps this species maintain condition even as cave energy levels
overall are declining — as judged by the decreasing condition of their counterparts in
the undisturbed cave end. However, as the pattern was only seen clearly in one of the
ten species we examined, caution is obviously warranted.
There is always a danger of overcollecting of cave fauna leading to reductions in
abundance, etc. (Culver, 1982). The collecting methods adopted by this study were
designed to minimise overcollecting. The rise in abundance during trip 3 shows that
these methods were successful.
8.1.5
Distance effects
The response of TA and fish to distance from the cave entrance was again different,
though in a different fashion than that seen for season. TA showed no clear response.
Fish show a clear response with rapidly decreasing abundance, richness and biodiversity with increasing distance from the cave entrance. Given that we believe all the
fish collected are epigean this is hardly surprising and simply represents a distance related decline from a source (the outside of the cave). We examined the response curves
of abundance and richness with distance against the typical (physical) “inverse-square”
model and the logarithmic decline suggested by Shaw and Davis (2000), like Shaw and
Davis (2000) we found that in most cases the log-based model provided better fits 4 .
However, the fit was far from good and the indication was that the response was likely
to be based on a non-linear parameterised model (not simply non linear in the sense
of not being of the form y = ax + b). However, given the limited number of distance
data we did not pursue this further.
The distance response shown by fish is particularly interesting as it may prove useful
as a management tool. Our data are insufficient to reliably test (insufficient power)
whether there is a difference in form between the disturbed and undisturbed response
4
though our examination of the data of (Shaw and Davis, 2000) found the inverse-square relationship gave better fits for their data!
70
Chapter 8. General discussion
71
curves5 . If further data was able to show a differential response this may allow fish
sampling to be used as an indicator of the distance disturbance effects intruding into a
cave.
8.1.6
Indicator species
ISA found several species of TA and fish whose presence was a significant indicator of
disturbed cave ends and, so by corollary, whose absence was an indicator of undisturbed
cave end. No species were found whose presence indicated undisturbed cave ends.
Only 1 species of TA was a significant indicator of disturbance. This was a beetle
of the family Staphilinidae. Two other species of TA proved to be marginal indicators
of disturbance.
Of the fish species identified as indicators of disturbed cave ends, both species (L.
birmanicus and R. paviei) are predators of aquatic invertebrates (Rainboth, 1996;
Smith, 1945). While we did not measure aquatic invertebrates this perhaps suggest
that disturbed cave ends may also have higher aquatic invertebrate abundance, probably
again as a result of energy enrichment or niche development.
Other results from the ISA showed that 7 species of fish were significant indicators
of distance into the cave. These species were found predominantly at the entrance and
exhibited a rapid decline in numbers with distance into the caves. With the exception
of Schistura magnifluvis, all belong to the family Cyprinidae, which feed on terrestrial
insects and zooplankton. The presence of light will be the strongest influence on the
location of these species. S. magnifluvis is benthic and usually found on gravel or
coarse sand feeding on organic detritus and debris.
The nature of ISA indicators may mean that they do not work well for cave species.
The species found in caves often display strong endemism and low abundance rates and
rare species are often not important in the ecosystem structure. However, rare species
are of great value for conservation and biodiversity value. It may be better to use levels
higher than species for ISA to avoid the problems caused by endemism.
8.1.7
Community composition
While all the biodiversity parameters we have considered to date are concerned with
α-biodiversity the CCAs give us an insight into biodiversity at the β – γ level.
Both TAs and fish generally show very clear within cave and within region similarities in composition, that is there is relatively little species turnover. The within cave
clustering is of course expected as caves are typically characterised by local faunas and
often endemics, though this is not always seen. Shaw and Davis (2000) reported that
the caves of Vancouver Island have many faunal elements shared with caves in Alaska.
In so far as the effects of disturbance on caves this is perhaps an encouraging sign as
often disturbance results in increased α biodiversity while resulting in reduced higher
level biodiversity. However, within regions, both TA and fish do show some weak
clustering according to disturbance level suggesting that β-biodiversity may also be
being affected by disturbance.
5
though the lack of any significant disturbance × distance interaction suggests there is no difference
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At the Order level for TAs it is interesting that Araneae — an Order pre-adapted
to cave life and the dominant taxa in our caves — and Orthoptera, dominated by
Rhaphidophoridae — a family also pre-adapted to cave life — show a bias toward
undisturbed cave ends.
The numerical dominance of a predatory taxa such as Araneae is interesting. The
dominance may be a result of our exclusion of micro-arthropods. It may also be a
result of the fact that the Order Araneae are remarkably consistently predators while
other Orders often consist of mixtures of predators, detritivores, etc. Certainly it would
be unusual to have a cave fauna community not dominated by detritivores, although
the high diversity of arachnids at all taxonomic levels in tropical caves has been noted
before (Deharveng and Bedos, 2000).
In his study of Batu Caves, Malaysia, McClure (1965) found that the fauna was
numerically dominated by flies, beetles and roaches. In terms of richness, the Diptera
were by far the most dominant (63 species). Coleoptera totalled 28 species and Araneae
were represented by just 9 species. This contradicts the findings of Deharveng and
Bedos (2000) and to a certain extent our own (total Diptera = 74 RTUs, Coleoptera
= 98 and Araneae = 84). The differences may be explained in the individuality of
biodiversity within caves. It may also be explained by different collecting techniques,
opportunities to identify species, etc. One thing is certain though, replicability is
essential when comparing cave biodiversity with human impacts.
8.1.8
Morphological modifications for hypogean life
Most of the TA collected and none of the fish showed any obvious morphological adaptations to cave life. Those TA that did possess adaptations have surface forms with
the same morphology, i.e. they are pre-adapted.
Although all our caves are also obviously continuous with epigean habitats as far as
TA are concerned the reduced motility and higher habitat and site fidelity typical of TA
means that adaptation and speciation of cave forms is more likely. However, none of
the TAs collected by us seem to be troglobites based upon their morphology. One group
that may be troglobites are flies of the family Keroplatidae, the larvae of which we have
never found outside of caves (although they do occur outside caves in Australia). Other
groups contain species that are likely to be troglophiles such as the Rhaphidophoridae
and various species of Diplopoda. Other groups are clearly “accidentals” with groups
such as Odonata being very unlikely to be anything other than accidental introductions
to caves.
All our caves were stream caves with mostly surface waters entering and passing
through the cave, Tham Tapan and Tham Thong being exceptions. In the case of Tham
Tapan the stream enters the cave passing through a hill (presumably an inaccessible
cave) and sinks underground at the other end, while Tham Thong enters on the surface
but sinks near the outflow end of the cave. This continuous connection with epigean
fish fauna, especially during the wet season, means that speciation of cave adapted
fish species seems unlikely and, indeed, all the fish collected were typical of torrential
streams in southern Thailand.
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Chapter 8. General discussion
8.1.9
73
Environment and physical
The measurements of co-factors such as entrance width, flood height, light penetration,
temperature and relative humidity, etc. do not suggest anything atypical from what
could be considered ’normal’ for caves of SE Asia (Smart, unpublished data). The fact
that the differences in these variables are small, both within and between the seven
study caves, substantiates this claim.
8.1.10
General fauna
Researchers have already described 8 species found within the caves of Phangnga
(Smart, unpublished data) including some remarkable relictual fauna (e.g. Dalens,
1989). These 8 species are also endemic to the area, in one case endemic to a single
pool of water (Dalens, 1989).
Our knowledge of Phangnga karst biodiversity is at a similar level to that of Maros.
The subterranean fauna is reasonably well known although many invertebrate species
remain undescribed. General observations of invertebrate fauna in Phangnga caves are
given by Deharveng and Bedos (2000), Deharveng and Bedos, (1988) and fish are listed
by Borowsky (1998). The 8 species descriptions completed to date are all invertebrates.
The most prominent finding is the remarkable Thailandoniscus annae (Dalens,
1989). This relictual species is the only Oniscidae crustacean adapted to an aquatic
habitat and the only one found in the Northern hemisphere. Other troglomorphic endemics are Eukonenia deleta, Siamacarus withi and Bogidiella thai. Cave-restricted
endemics are Phricotelphusa deharvengi and Ptomaphaginus leclerci. Other endemics
are Paranura globulifer , P. leclerci and Annina fustis.
Overall, to date, there have been at least 140 species described from caves in Thailand (Smart, unpublished data). This number of species has resulted from remarkably
little research and it seems likely that there are many new species waiting to be discovered and described.
There has never been a systematic survey of the biodiversity of any cave in southern
Thailand, or any attempt to link cave biodiversity with human impacts.
TA
It is difficult to discuss the TA fauna collected by this study as they have not been
identified to species level. However, the following points can be highlighted.
It seems likely that some of the specimens collected represent new, undescribed
species, although we will never know without further taxonomic work. This work will
also remove those RTUs that are double (or more) counted due to immature/mature
or male/female differences.
Three separate communities of TA can be distinguished by community composition
analysis (figure 7.9). The three groups are based, as expected, on region and separate
into the Satun group, Phangnga group and Chumphon group, although the disturbed
end of Tham Tapan plots some distance away from the main Phangnga cluster.
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Fish
The spiny eel of the genus Macrognathus sp. is an undescribed new species while the
status of Lepidocephalichthys sp. and Hemimyzon sp. is still uncertain.
Nearly all species found were typical for torrential streams throughout southern
Thailand, with only six species being regarded as exceptions. Two exceptions were
Opsarias bernaski (Coumans) and Poropuntius melanotaenia (Roberts) which are restricted to the Tenasserim zone. The other four species are non-native species (or
hybrids) — hybrid catfish: Clarias gariepenus × C. macrocephalus; Nile Tilapia: Oreochromis niloticus (Linnaeus); South American Swordtail: Xiphophorus helleri; and
Mosquito fish: Gambusia affinis (Baird & Girald). These four species were only found
in Tham Phung Chang. It is likely that they have been released as part of Buddhist
merit making practices.
Despite the very typical nature of the fish fauna it is still possible to clearly separate
three communities based upon community composition (Figure 7.19). First, the Satun
group (JKT and KKL) with larger streams containing Rasbora bangkanensis (Bleeker),
Glyptothorax spp. and plenty of Silurid catfishes as the dominant taxa. Second, the
Phangnga group (NM1, PCH, TPN and TNG) with small streams hosting loaches
(Lepidocephalichthys spp. and Schistura magnifluvis (Kotellat). These species typically
inhabit torrential streams with gravel or sand beds and they are also indicative of S
and E Thailand. Lastly, the Chumphon group, in fact only one cave (NLY). This cave
contains a large and rapid torrential stream. The fish are rather different from the
other caves in having Tenasserim fish fauna. Indicator species are O. c.f. puchellus
and P. melanotaenia.
There are 2 further species of interest: Aplocheilus panchax (Hamilton) and Glossogobius aureus (Akihoto and Meguro). These species are normally found in brackish
water and their presence indicates that the stream they inhabit is connected to mangrove forest. Also, the relatively high salinity of karst water allows these fish to swim
far upstream of their normal habitats.
8.1.11
Limitations
We could not find any non-disturbed caves in which to conduct our study as mentioned
earlier and all of our undisturbed caves were relatively so. It would have been better if
we could have acquired non-disturbed caves for the study as this would have made it
possible to compare between original cave biodiversity and impacted cave biodiversity.
As it is, we are able only to compare between less impacted biodiversity and more
impacted biodiversity.
Another limitation of this study is the fact that only macro-TAs and fish were
looked at. Other groups, such as micro-arthropods, non-fish aquatic fauna, etc. were
not investigated. We are confident though that the groups studied do represent cave
biodiversity as a whole and that the changes we have noted in these two groups would
also be observed in others.
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Chapter 8. General discussion
8.2
75
The whole
The hole is indeed more than the sum of the parts! Caves are certainly more than
holes in rocks and the biodiversity of caves and the effects of disturbance upon them are
certainly more than we have been able to examine in this study. This work has simply
added another piece to the puzzle. Hopefully, the results of this study will provide
scientists and managers with valuable data that will be of direct use in furthering
understanding and improving conservation.
Perhaps the most important point to come out of this work is that more biodiversity
does not necessarily equal better biodiversity. We have shown that disturbance of caves
increases biodiversity and this fact should not be used as congratulatory, but rather
an alert. Animal populations within most caves are extremely small, and many cave
species are on the verge of extinction (Moore and Sullivan, 1997). Due to their low
abundance rates and high degrees of endemicity, cave fauna should to be regarded as
a conservation priority. It is essential that everything possible is done to protect them
from energy enrichment, invasive species and other threats.
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Chapter 9
Conclusions and recommendations
He who adds not to his learning
diminishes it.
The Talmud
. . . knowledge of nature is an
expanding universe, continually
creating ever greater circumstances
of ignorance, a concept that can be
expressed in the words: “The more
we know, the less we know.”
Chargaff 1969
9.1
Conclusions
This study has shown that disturbance due to human visitation to stream caves in
southern Thailand has caused changes in the biodiversity of caves in terms of richness
and abundance. People are able to change the cave habitat in various ways, both
intentionally and accidentally and we hope that cave managers will take notice of
these findings. The most important task now is to spread the message to the general
public and managers about the uniqueness of cave fauna and the problems that people
can cause. We hope that the results will not be misinterpreted in that a greater
biodiversity is equal to a better biodiversity and that alien species are kept out of cave
ecosystems. Cave animals are often few in number and highly endemic. Many of them
are endangered and need proper conservation strategies to ensure their survival.
9.2
Management recommendations
1. Extra caution with regard to development of high energy caves: Given
the clear evidence presented here showing large effects on cave biodiversity as a
result of human disturbance, it seems wise to re-evaluate the belief that “highenergy caves are suitable for development”.
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2. Discouragement of merit making releasing of exotic fauna: Thailand
has a relatively rich troglobitic fish fauna and it would be a considerable and
ironic tragedy if this fauna was endangered as a result of “merit-making” fish
releases. There should be greater education and enforcement measures aimed at
stopping the practice of releasing animals into non-native habitats. Such measures
would be in accord with Thailand’s commitment to better control invasive alien
species which are now regarded by IUCN as the second most important cause of
biodiversity loss.
3. Better control over visitors: cave managers need to exercise better control
over the actions of visitors in general. Things like dropping litter, writing graffiti,
smoking and walking off the footpath (if present) are widespread and need to be
stopped.
4. Use management strategies known to reduce impacts: the use of nonbiodegradable materials in cave infrastructure should be encouraged and the
time that lighting systems are switched on should be reduced. We also follow
the view of Watson et al. (1997) that already impacted caves should be restored
and developed for tourism rather than the opening of new caves.
9.3
Research recommendations
The following recommendations come directly from this study and from meetings between cave and karst researchers in Bangkok and in Phuket.
1. Regional level confirmation study: although the results of this study are
clear, the level of replication (seven caves in southern Thailand) means that
caution should be exercised in generalising too broadly across the region or more.
Similar rigorous work needs to be conducted in other areas.
2. Elucidation of mechanisms: while we speculate on mechanisms in this report,
particularly the idea of the “Paradox of Enrichment” further studies are needed
to clearly demonstrate the mechanism involved. Such studies would not only
provide valuable scientific information but also be important for management as
it may be that strategies can be applied effectively to mitigate impacts.
3. Need more work on the high energy caves paradigm: it would be useful
to conduct a similar study on low energy caves to confirm that high energy caves
are no less susceptible to disturbance.
4. Need taxonomic support for cave TAs: this study identified all TA species
as RTUs. The lowest level where a name could be applied was genus level for a
few, certain groups, e.g. the Amblypygids. Most groups could not be identified
to this level. Work and support is most certainly needed in this area.
5. Other cave taxa remain little known: this study looked at fish and neglected
all other aquatic fauna. Micro-invertebrates were also left out. These taxa, and
others not studied, must be regarded as a research priority.
78
Chapter 10
Acknowledgements
Thanks to ARCBC, ASEAN and the EU for funding and so making this research possible.
We would first like to thank Martin Ole Odderskov (Zoological Museum, University of Copenhagen — Coleoptera), Jan Peterson (Zoological Museum, University of
Copenhagen — Coleoptera), Sadahiro Ohmomo (JIRCA, Thailand — Coleoptera),
Peter Weygoldt (Institut für Biologie I (Zoologie), Albert-Ludwigs-Universität — Amblypygi), Kazuo Ogata (Institute of Tropical Agriculture, Kyushu University — Formicidae), Korakot Damrak (KU, Thailand — Staphylinidae and Trichoptera) and Jamnongjit Phasook (KU, Thailand — Psycodidae) for their assistance with identification
of specimens. Thanks also to Phanwadee Thamrongwang (DNPW) for arranging analysis of our water samples.
Thanks to Chet Phuangjit (Chief of Ao Phangnga National Park), Yotchai Boonyanet
(Chief of Khao Banthat Wildlife Sanctuary), Wirote Phuangphakhisiri (Chief of Typhoon Gaye Rehabilitation Project), and Anan Charoensuk (Chief of Namtok Ngao
National Park) for providing us with accommodation during field work.
Sriwieng Chuayluer and Prasert Chuayluer provided us with good food and good
company while in the field at Satun and thanks for their help during a medical emergency during our third trip.
We are grateful to Chutima Dokmai, Sirada Sommin, Nalin U-glud, Saichon Duangloy, Pranee Srirabai, Banjongluck Wangsilabut for assistance in the laboratory and
preparation for the field trip. We are also grateful for the efforts of Chris Dickinson
who was involved in drafting the original research proposal.
Thanks to Sahakit Phakdee quarrying company for granting access permission
to Tham Phung Chang. Thanks to Weecheep Jaturabundit (Provincial Forester—
Phangnga) and Suwat Narongrid (Assistant District Forester—Phangnga) for assisting
us to gain permission to access Tham Phung Chang.
Thanks to the DNPW for granting permission to collect in protected areas.
Thanks to A. Bedos, E. Cholik, L. Deharveng, R. Marwoto, C. Rahmadi, Y. Suhardjono and A. Suyanto the lecturers of the training course in Indonesia on “Invertebrates
taxonomy with special reference to less well-known groups”. This course was funded
by ARCBC and of our project staff, Patpimon Sawai, attended.
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80
Part III
Appendices
81
Appendix A
Publications and Presentations
(developed as part of the project)
During the course of the project several reports and presentations have been prepared
and presented–with the hope that more will follow.
A.1
Technical and administrative reports
1. Anon. (2002) “The Effects of Human Impacts on Cave and Karst Biodiversity:
Thailand Component: Progress report for the period: February–July 2002” submitted to ARCBC.
2. Anon. (2003) “The Effects of Human Impacts on Cave and Karst Biodiversity:
Thailand Component: Annual Report Year 1” submitted to ARCBC.
3. Cunningham, RJ (2001) A progress report for the project “The Effects of Human
Impacts on Cave and Karst Biodiversity: Thailand Component” submitted to
National Research Council of Thailand (NRCT).
4. Cunningham, RJ (2002) A progress report for the project “The Effects of Human
Impacts on Cave and Karst Biodiversity: Thailand Component” submitted to
NRCT.
5. Sawai, P (2002) “Field trip report: 12-28 August 2002 at Maros, Indonesia”,
submitted to ARCBC.
6. Smart, D (2001) A progress report for the project “The Effects of Human Impacts
on Cave and Karst Biodiversity: Thailand Component” submitted to the NRCT.
7. Smart, D (2002) A progress report for the project “The Effects of Human Impacts
on Cave and Karst Biodiversity: Thailand Component” submitted to the NRCT.
8. Smart, D (2003) A progress report for the project “The Effects of Human Impacts
on Cave and Karst Biodiversity: Thailand Component” submitted to the NRCT.
9. Smart, D and Cunningham, RJ (2001) “Field trip report: Phangnga caves” a
report submitted to the Cave Management Committee, Natural Resources Conservation Office, Royal Forest Department.
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10. Smart, D and Cunningham, RJ (2002) “Field trip report: assorted caves of southern Thailand” a report submitted to the Forest Environment Research and Development Division, Forest Research Office, Royal Forest Department.
A.2
Presentations at scientific meetings
In addition to these reports results from the project have been presented at two scientific
workshops.
A.2.1
Workshop on “The effects of human impacts on cave and
karst biodiversity”
During 25-28 May 2003 a workshop was held at Hat Nai Yang, Phuket, Thailand. The
participants of the workshop were Yayuk Suhardjono, Ristiyanti Marwoto and Cahyo
Rahmadi from our sister project (The Effects of Human Impacts on Cave and Karst
Biodiversity: Indonesian Component), Aida Lapis (ARCBC, Philippines) and the entire
Thai team1 . The goal of the workshop was to allow the Indonesian and Thai researchers
an opportunity to discuss their research project progress and results obtained to date
as well as have discussions on means to improve regional cave and karst research and
management in the future. The Thai team presented the following presentations during
this workshop.
1. Cunningham, RJ (2003) “Experimental design and sampling procedures”
2. Cunningham, RJ (2003) “Early results of the effects of human disturbance on
non-Araneae terrestrial arthropods”
3. Hutacharern, C (2003) “An introduction to the project ‘The Effects of Human
Impacts on Cave and Karst Biodiversity: Thailand Component’ its background
and objectives”
4. Janekitkarn, S (2003) “Preliminary results of the effects of human disturbance on
cave fish fauna”
5. Smart, D (2003) “An introduction to caves and cave management in Thailand”
6. Smart, D (2003) “An introduction to our study caves”
7. Vichitbandha, Patchanee (2003) “Preliminary results of the effects of human disturbance on Araneae”
A.2.2
ARCBC Regional Research Grant Conference: Building bridges
between ASEAN and EU researches[sic]
The Regional Research Grant Conference, held in Bangkok during December 1-4 2003,
was the gathering of researchers from the projects funded by ARCBC and EU specialists from various institutions. As part of this conference Chaweewan Hutacharern, as
1
in addition management staff from the Department of National Parks, Wildlife and Plant Conservation were invited, however, no one attended
84
Appendix A. Publications and Presentations
85
project leader, also gave a presentation entitled: “The effects of human impacts on cave
and karst biodiversity: Thailand component” which presented the results of the project
with regard to fish biodiversity. Researchers from our project also joined the discussion
and presentation of the Theme “Cave and Karst Biodiversity” (ARCBC, 2004).
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86
Appendix B
Taxonomic checklist
Table B.1 provides a taxonomic checklist of the fish collected during the project. Of the
three species lacking specific epithets Macrognathus sp. is an undescribed new species
while the status of Hemimyzon sp. and Lepidocephalichthys sp. is still uncertain.
No similar list is available for TA as the vast majority of material is unidentified
and only identified by voucher codes.
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Table B.1: Checklist of fish species collected during the study (includes trips 1-3)
disturbance levela
disturbed undisturbed
Family
Binomial
1
Ambasseidae
Pseudambassis siamensis (Blyth)
2
3
4
5
Andrianichthyidae
Bagridae
Bagridae
Bagridae
Aplocheilus panchax (Hamilton)
Batasio tengana (Hamilton)
Hemibagrus filamentosus (Fang & Chaux)
Leiocassis siamensis (Regan)
6
6
Bagridae
Bagridae
Mystus nemurus (Valenciennes)
Mystus singaringan (Bleeker)
8
Balitoridae
Hemimyzon sp.
9
10
Balitoridae
Balitoridae
Homaloptera leonardi (Hora)
Homaloptera smithi (Hora)
11
12
13
14
Balitoridae
Belontiidae
Channidae
Channidae
Schistura magnifluvis (Kotellat)
Trichogaster trichopterus (Pallas)
Channa gachua (Hamilton)
Channa lucius (Cuvier)
15
Channidae
Channa striata (Bloch)
16
Cichlidae
Oreochromis niloticus (Linnaeus)
17
Clariidae
Clarias gariepenus × C. macrocephalus HYBRID
18
Clariidae
Clarias teysmanni (Bleeker)
19
20
21
Cobitidae
Cobitidae
Cobitidae
Lepidocephalichthys birmanicus (Rendhal)
Lepidocephalichthys hasselti (Valenciennes)
Lepidocephalichthys sp.
22
Cyprinidae
Brachydanio albolineatus (Blyth)
23
24
25
26
27
Cyprinidae
Cyprinidae
Cyprinidae
Cyprinidae
Cyprinidae
Brachydanio kerri (Smith)
Cyclochilichthys apogon (Valenciennes)
Danio aequipinnata (McClelland)
Danio regina (Fowler)
Esomus metallicus (Ahl)
28
Cyprinidae
Hampala macrolepidota (Valenciennes)
29
30
31
Cyprinidae
Cyprinidae
Cyprinidae
Mystacoleucus marginatus (Valenciennes)
Neolissochilus blanchi (Pellegrin and Fang)
Neolissochilus soroides (Duncker)
32
33
34
Cyprinidae
Cyprinidae
Cyprinidae
Neolissochilus stracheyi (Day)
Opsarias puchellus (Smith)
Opsarias bernaski (Couman)
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
Continued following page. . .
: species absent; ~: species present but only one individual; other sizes represent relative
abundance (log2 scale)
ax
0
88
89
Appendix B. Taxonomic checklist
Table B.1: continued
Family
Binomial
35
Cyprinidae
Osteochilus hasselti (Valenciennes)
36
Cyprinidae
Osteochilus microcephalus (Valenciennes)
37
38
39
40
Cyprinidae
Cyprinidae
Cyprinidae
Cyprinidae
Poropuntius melanotaenia (Roberts)
Rasbora bangkanensis (Bleeker)
Rasbora daniconius (Hamilton)
Rasbora dusonensis (Bleeker)
41
42
43
44
Cyprinidae
Cyprinidae
Cyprinidae
Gobiidae
Rasbora paviei (Tirant)
Systomus binotatus (Valenciennes)
Systomus lateristriga (Valenciennes)
Glossogobius aureus (Akihoto and Meguro)
45
Gobiidae
Pseudogobiopsis siamensis (Fowler)
46
Hemiramphidae
Dermogenys pusilla (van Hasselt)
47
Mastacembelidae
Macrognathus sp.
48
49
Mastacembelidae
Poecilidae
Mastacembelus favus (Hora)
Gambusia affinis (Baird & Girald)
50
Poecilidae
Xiphophorus helleri (Heckel)
51
Siluridae
Ompok bimaculatus (Bloch)
52
53
55
55
Siluridae
Siluridae
Sisoridae
Sisoridae
Pterocryptis torrentis (Kobayakawa)
Silurichthys schneideri (Volz)
Glyptothorax fuscus (Fowler)
Glyptothorax major (Boulenger)
56
Sisoridae
Glyptothorax trilineatus (Blyth)
57
Synbranchidae
Monopterus albus (Zuiew)
89
disturbance level
disturbed undisturbed
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
~
u
~
~
~
~
~
~
~
~
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90
Appendix C
Location, cave and station maps and
sketches
Maps showing cave locations, detailed maps of caves and sketch maps of sampling
stations arranged North-South according to groups (based broadly on amphur).
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C.1
Chumphon
One cave, Tham Than Nam Lot Yai, was located in amphur Sawi in Chumphon
province. A brief description of the cave, a detailed location map, a cave plan and
station sketches are provided below.
C.1.1
Tham Than Nam Lot Yai [Figures C.2 and C.3]
This is a very large river cave. It passes through a 5 km long ridge (Khao Talu) nearly
500 m in height. Powerful winds blow through the cave.
Disturbed (inflow) [Figure D.1(c), page 120]
Located 1 km from a main road intersection at the end of a short unsurfaced road. This
end of the cave is well signposted and there is a car park and toilet facilities. Massive
amounts of flood debris characterise this entrance, which is almost completely choked
with large tree trunks, logs, bamboo and other assorted junk. A small, seasonal passage
to the left bypasses the debris and soon meets the river again at a series of rapids in a
large boulder strewn passage. From this point onwards, the passage increases in size to
30 m wide by 10 m high. The gradient decreases and the river becomes more sedentary
flowing through wide, deep pools. Thick mud covers the floor. Once a year, local school
children are recruited to cleanup the entrance area in anticipation of the hundreds of
tourists that visit during the Songkhran (Thai New Year) festival in April.
Undisturbed (outflow)
A well-maintained dirt road passes within 100 m of the entrance. A small track turns
off this and leads to the entrance, which has a wooden ladder descending down boulders
to river level. The entrance is very large, 30 m wide by 8 m high, with extensive gravel
and mud banks. The river flows swiftly in a channel against the right wall. Inside, the
cave increases in size and at one point becomes almost 70 m wide. A primitive, and
probably dangerous, electric light system has been installed at this end of the cave,
though it doesn’t appear to be used very often if it even works.
92
Appendix C. Location, cave and station maps and sketches
93
Figure C.1: Location of Tham Than Nam Lot Yai in amphur Sawi, Chumphon [based
upon 1:50 000 topographic map sheet 4729 II]
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Figure C.2: Tham Than Nam Lot Yai floor plan (Source: Deharveng and Bedos
(1988))
94
Appendix C. Location, cave and station maps and sketches
95
Figure C.3: Sketches of sampling stations in Tham Than Nam Lot Yai (legend on
page 117)
95
96
C.2
The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Phangnga (Amphur Mueang)
Two caves, Tham Phung Chang and Tham Tapan, were located in amphur Mueang
in Phangnga province. A brief description of the caves, a detailed location map, cave
plans and station sketches are provided below.
Figure C.4: Locations of Tham Phung Chang and Tham Tapan in amphur Mueang,
Phangnga [based upon 1:50 000 topographic map sheet 4725 IV]
96
Appendix C. Location, cave and station maps and sketches
C.2.1
97
Tham Phung Chang [Figures C.5, C.6]
A single stream passage cave passing through the base of Khao Luk Chang. Quite
pretty with numerous interesting features and popular with tourists.
Disturbed (outflow)
Located behind a temple in Phangnga town and very well signposted. There is a car
park and restaurants. A dam at the entrance has flooded the floor at this end of the
cave. The water is generally waist deep with a muddy bottom. One short section
requires swimming. At about 180 m from the entrance a skylight enters on the right.
Tourists are taken into this end of the cave on inflatable boats. About 250 m inside, a
rock bridge is met where the tourists have to get out and climb over using permanently
installed wooden steps. On the other side they continue their way using bamboo rafts.
Undisturbed (inflow) Although it is possible to drive to within 100 m of this
entrance, the track is privately owned and public access is not normally granted. The
large, multiple entrances of this end quickly close down to a small passage 2 m in
diameter. Some 100 m inside this opens up into a chamber floored with boulders and
flowstone and a reasonable amount of bat guano. Wooden ladders climb up here to
a dry, upper level bypassing a very low crawl in the stream. The upper level quickly
drops back down to the stream, which flows on into a wide, low passage with a gravel
floor.
97
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Figure C.5: Tham Phung Chang floor plan (Source: Maffre et al. (1986))
98
Appendix C. Location, cave and station maps and sketches
99
Figure C.6: Sketches of sampling stations in Tham Phung Chang (legend on page 117)
99
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
C.2.2
Tham Tapan [Figures C.7, C.8]
An attractive cave with a variety of features. Consists of two stream inlet passages
draining separate valleys on the surface. These connect underground to form a single,
large outlet passage.
Disturbed (outflow) [Figure D.1(a), page 120]
Located in Phangnga town within the grounds of a large temple, which has a car park
and restaurants and is very well signposted. This end of the cave is fully developed
with electric lighting, concrete paths and bridges and is heavily visited by tourists.
The passage is large and walking is easy, though very muddy away from the footpath.
Beyond the developed section, the stream becomes gravel floored and bats provide
guano. However, artificial light from the fluorescent tubes still reaches this point.
Undisturbed (inflow)
From the temple at the outflow end, follow a steep track up and around the other side
of the mountain and then walk for 30 minutes through overgrown forest. The entrance
to this end is a large collapse doline requiring a steep scramble down boulders to gain
access to the stream. This is met at the lowest point in a low, wet crawl. A varied
passage follows with alternating easy walking and low crawling over gravel. A small
waterfall and deep plunge pool add further interest.
100
Appendix C. Location, cave and station maps and sketches
Figure C.7: Tham Tapan floor plan (Source: Kiernan (1986))
101
101
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Figure C.8: Sketches of sampling stations in Tham Tapan (legend on page 117)
102
Appendix C. Location, cave and station maps and sketches
C.3
103
Phangnga (Amphur Thap Phut)
Two caves, Tham Thong and Tham Nam 1, were located in amphur Thap Phut in
Phangnga province. A brief description of the caves, a detailed location map, cave
plans and station sketches are provided below.
Figure C.9: Location of Tham Thong and Tham Nam 1 in amphur Thap Phut,
Phangnga [based upon 1:50 000 topographic map sheet 4726 III]
103
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
C.3.1
Tham Thong/Tham Lot [Figures C.10, C.11]
An interesting cave carrying two, equally tiny streams that join not far inside the inflow
end. Local people use the cave as a thoroughfare through the mountain. It has two
names because the people living at the inflow end call it Tham Thong and the people
at the other end call it Tham Lot.
Disturbed (inflow)
An unsurfaced, though decent road, passes within 20 m of the entrance. Heading in
from the large, compound entrance is a spacious passage 9 m wide by 5 m high. After
50 m a side passage joins on the right carrying a small inlet. Just beyond here a
prominent flowstone cascade is reached. This is either climbed over or bypassed by a
high level passage to the left. Beneath the flowstone is a deep pool with a water pipe
feeding out to the cave entrance. After this point, the cave gets smaller in size and
starts to meander more with gravel banks and short riffles.
Undisturbed (outflow)
After driving 2 km on a rough track through a rubber plantation, walk 1 km with no
real footpath. The stream flows out of this entrance only during floods. In dry weather
it disappears down a small hole about 50 m inside. Beyond this point though, the water
flow is perennial. The passage at this end of the cave is variable in size, being quite
large in places and small in others. At one point it reduces to a low crawl over gravel.
104
Appendix C. Location, cave and station maps and sketches
Figure C.10: Tham Thong floor plan (Source: Maffre et al. (1986))
105
105
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Figure C.11: Sketches of sampling stations in Tham Thong (legend on page 117)
106
Appendix C. Location, cave and station maps and sketches
C.3.2
107
Tham Nam 1 [Figures C.12, C.13]
The stream passage of this cave is around 550 m long and passes through a ridge. Few
people visit the cave outside of local people catching fish and bats for food. Blasting
at a nearby quarry provides additional amusement.
Disturbed (inflow)
Walk 15 minutes through a rubber plantation on a reasonably well-trodden footpath.
This entrance is spacious with easy walking over gravel banks and a meandering stream
with pools and riffles. However, a low section is soon reached that requires stooping
through a waist deep pool. After this, the passage enlarges again and continues fairly
straight for 60 m to a bend with a deep pool. Following this is a chamber with a gravel
and flowstone floor. A fair number of bats roost here.
Undisturbed (outflow)
Walk 30 minutes through fields following farmers’ footpaths. This entrance has been
dammed using sand bags to act as a local water supply, though the ponded water does
not extend very far into the cave. This end of the cave starts as a long, straight rift
passage with the stream flowing along a narrow trench. Progress is made by traversing
along bare rock ledges on the left wall. After 120 m the passage turns a corner and
becomes wider with a gravel floor. A fair-sized bat colony roosts at the furthest point
in.
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Figure C.12: Tham Nam 1 floor plan (Source: Deharveng and Bedos (1988))
108
Appendix C. Location, cave and station maps and sketches
109
Figure C.13: Sketches of sampling stations in Tham Nam 1 (legend on page 117)
109
110
C.4
The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Satun
Tham Jet Khot is located in amphur La-ngu while Tham Khong Kha Lot is actually in
amphur Pa Bon, Phattalung province but it is accessed from La-ngu and so we include
it in Satun. A brief description of these caves, a detailed location map, cave plans and
station sketches are provided below.
Figure C.14: Location of Tham Jet Khot, amphur La-ngu, Satun province, and Tham
Khong Kha Lot, amphur Pa Bon, Phattalung province [based upon
1:50 000 topographic map sheet 4923 II]
110
Appendix C. Location, cave and station maps and sketches
C.4.1
111
Tham Jet Khot [Figures C.15, C.16]
This is a very large, railway tunnel-type cave taking a small river through a low ridge.
The passage averages 15 m wide by 30 m high and meanders gently around long, sweeping bends. It floods severely in wet weather.
Disturbed (outflow)
Even though a long drive on increasingly worse roads and tracks is necessary to reach
this end, it is well signposted and has a small “resort” with a cleared area used for car
parking. From here it is a short walk upriver to a massive entrance. Progress is made
into this end by walking over extensive gravel banks and wading across shallow sections
of the river. Deep pools occur on the outside of the bends. There is a skylight high in
the roof about 250 m in and a large bat colony. This end is a popular party venue with
local people. Occasionally, organised tour groups are taken in by boat.
Undisturbed (inflow) [see page ii]
Access to this end was originally a long rough track (20 km) followed by walk through
the forest on a footpath. During the timeframe of this study, construction work on a
wide, new tarmac road and car park was being carried out and was nearly finished by
trip 3. At the cave entrance, the river flows swiftly between boulders. Immediately
inside, the water enters a slower-moving pool filling the width of the passage. Wading
and swimming across this leads to large gravel banks with shallow riffles in between.
111
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Figure C.15: Tham Jet Khot floor plan (previously unpublished)
112
Appendix C. Location, cave and station maps and sketches
113
Figure C.16: Sketches of sampling stations in Tham Jet Khot (legend on page 117)
113
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
C.4.2
Tham Khong Kha Lot [Figures C.17, C.18]
A highly attractive cave retaining many natural qualities due to its remote location.
The single stream passage connects via a vertical rift with the overlying Tham Phu
Pha Phet (Gray, 2001). Flood debris is stuck to the roof and walls throughout the
cave indicating very severe flooding in the rainy season.
Disturbed (inflow)
Located behind the temple at Tham Phu Pha Phet, which is signposted and has car
parking and tourist facilities. Work on widening and surfacing the access road was being
carried out as this study proceeded. Inside the 10 m-wide, stooping sized entrance,
the passage quickly enlarges to walking sized proportions with meandering bends and
extensive gravel banks. Nicely decorated with speleothems.
Undisturbed (outflow)
At the end of a very long drive on ever worsening tracks followed by a walk of 90
minutes through the forest and down a river valley. This end is very remote and rarely
visited. Beyond the complex, interconnected passages of the entrance, the cave narrows
to a single straight rift. Part way along the rift, the water deepens at an overhanging
flowstone necessitating a short swim. Continuing past this, the water shallows again
and forms riffles over gravel and bare rock. At the end of the rift, the passage bends
to the right and changes to a wide, meandering canyon with gravel banks.
114
Appendix C. Location, cave and station maps and sketches
Figure C.17: Tham Khong Kha Lot (Source: Smart and Cunningham (2002))
115
115
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
Figure C.18: Sketches of sampling stations in Tham Khong Kha Lot (legend on
page 117)
116
Appendix C. Location, cave and station maps and sketches
Passage wall
Stalactite
Passage wall-inaccessible
Stalagmite
Passage wall-flowstone
Column
Vertical step
Flowstone
Stream flow direction
Gour pools
Boulder
Coralloids
Gravel
Guano
Sand
Green plant
Mud
Tree
Bare limestone
Root
Still or slow-moving water
Slope indicator
Flowing water
Concrete
Rapids
Wooden steps
Flood debris
Buddha statue
Figure C.19: Legend for station maps
117
117
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
118
Appendix D
Contact sheet
A selection of project photographs showing locations, activities or animals related to
the project.
see next page
119
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The Effects Of Human Impacts On Cave & Karst Biodiversity: Thailand Component — Final Report
(a) A metal bridge and flourescent lights in
Tham Tapan, an obvious example of how caves
can be disturbed by human activity
(b) A wooden bridge in Tham Phung Chang.
While this bridge is “biodegradeable” that very
fact means that the bridge represents an unnatural source of energy in this cave — another type
of disturbance
(c) Tham Than Nam Lot Yai (disturbed entrance) in flood, the seasonal flooding of these
stream caves is a very important source of energy and natural disturbance for these caves
(d) Sampling for terrestrial arthropods using a
quadrat and manual searching
(e) Weighing and measuring of fish prior to release
(f ) Opsarias bernaski was collected from Tham
Than Nam Lot Yai, this being the first record of
this species from Thailand
Figure D.1: A selection of project photographs — set 1
120
Appendix D. Contact sheet
121
(a) Participants in the workshop “The effects of human impacts on cave and karst biodiversity”.
Front row (l-r): Aida Lapis (Philippines), Ristiyanti Marwoto (Indonesia), Yayuk Suhardjono (Indonesia), Chaweewan Hutacharern (Thailand), Patpimon Sawai (Thailand), Phuwadol Vichitbandha
(Thailand), Patchanee Vichitbandha (Thailand), Ruenchit Phukthair (Thailand); Second row (l-r):
Sommai Janekitkarn (Thailand), Ophart Chamason (Thailand), Cahyo Rahmadi (Indonesia), Watana
Sakchoowong (Thailand), Prasong Hemapan (Thailand), Dean Smart (Thailand); Missing: Robert
Cunningham (Thailand)
Figure D.2: A selection of project photographs — set 2
121
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122
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