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... FOREST OF SINGAPORE Introduction The main island of Singapore is located off the southern tip of the Malay Peninsula and most of the island consisted of tropical lowland rain forest for much of the. .. demonstrated this pattern of functional group succession by conducting a study of the ant community within a secondary tropical rain forest in Singapore Using the mathematical models, the ages of. .. New Caledonia Barro Colorado Island, Panama Barro Colorado Island, Panama Atherton Tablelands, Australia Kununurra, Australia Vicosa, Brazil Mkomazi Game Reserve, Tanzania Popondetta, Papua New
ANT FUNCTIONAL GROUP SUCCESSION DYNAMICS CORRELATES WITH THE AGE OF VEGETATION SUCCESSION: DATA ANALYSIS OF WORLDWIDE STUDIES AND A CASE STUDY OF A SECONDARY TROPICAL RAIN FOREST IN SINGAPORE THAM KOON HUNG, ANDREW (B.Sc (Hons.), NUS) A THESIS SUBMITTED FOR THE DEGREE OF MASTER OF SCIENCE DEPARTMENT OF BIOLOGICAL SCIENCES NATIONAL UNIVERSITY OF SINGAPORE 2006 a ACKNOWLEDGEMENTS My deepest gratitude to Associate Professor Li Daiqin, who not only guided me and kept me focused on this project but also allowed me the freedom to think and play with ideas I will look back many years from now and wonder where I would have gone without him My heartfelt thanks to members of the Spider Lab, whose friendship kept me young-at-heart, especially Jeremy Woon for all the “lunge” sessions, Matthew Lim for the “tech support”, Olivia Tan for evaluating the standards of my jokes, Seah Wee Khee for actually laughing at them, Chris Koh for the entertaining CDs I thank Reuben Clements and Kelvin Peh for their patience and enormous help with data analysis, Darren Yeo for his initial advice on being an ant taxonomist, the National Parks Board, Chew Ping Ting and Benjamin Lee for approving the research permit, supplying the GIS data and guidance when I was lost in the jungle To Faith, my precious daughter, for giving me the impetus to finish this thesis To my ever-lovely, then-girlfriend and now wife, Geraldine, without whom I may never have had the belief in myself to anything meaningful in life Finally, to JC, my constant but silent friend: thanks for forgiving my nonsense “Go to the ant, thou sluggard; consider her ways, and be wise ” - The Bible i TABLE OF CONTENTS Page ACKNOWLEDGEMENTS i TABLE OF CONTENTS ii ABSTRACT iii LIST OF TABLES iv LIST OF FIGURES v CHAPTER GENERAL INTRODUCTION CHAPTER ANALYZING PUBLISHED DATA TO STUDY THE 10 SUCCESSION ECOLOGY OF ANTS CHAPTER Introduction 10 Material and Methods 15 Results 20 Discussion 33 ANT SUCCESSION DYNAMICS IN A SECONDARY 41 RAIN FOREST OF SINGAPORE CHAPTER Introduction 41 Material and Methods 46 Results 55 Discussion 66 GENERAL DISCUSSION 72 REFERENCES 81 APPENDIX 95 ii ABSTRACT Many studies have shown that ant communities respond to changes in the environment but some questions remain unanswered What functional groups dominate different types of ecosystems? What groups succeed one another? Are there trends in the succession of ant communities in relation to vegetation succession? To answer these questions, data analysis of existing studies from six terrestrial ecosystems, viz tropical rain forest, montane forest, temperate forest, desert, subtropical grassland, tropical bushland, was performed I showed that each habitat had different dominant functional group In tropical rain forests, Opportunist and Tropical Climate Specialist functional groups were the pioneer community that established in young, disturbed vegetation These two groups were succeeded by Cryptic Species, Generalized Myrmicinae and Specialized Predator groups during vegetation succession Sigmoidal mathematical models best described the decline and growth of these two communities, respectively I demonstrated this pattern of functional group succession by conducting a study of the ant community within a secondary tropical rain forest in Singapore Using the mathematical models, the ages of different locations in this forest were estimated This pattern of functional group succession implies that ants may be used as bioindicators during forest rehabilitation Directed studies on specific groups may be conducted to assess their impact on forests at different stages of succession The mathematical models represented a first step in developing tools for estimating forest age where historical records are lacking iii LIST OF TABLES Page Table Summary table of studies on ants 19 Table Description of the typical vegetation found in each of the four Forest 52 Types in Singapore Table Study site, forest type and environmental variables 54 Table Variance in species data (r2) represented by the three axes of 62 ordination and the coefficients of correlation of significant environmental variables with axes two and three Table MRPP Results of average within-group (Forest Type) distances and 63 pair-wise comparisons between groups iv LIST OF FIGURES Page Figure Proportions of functional groups in tropical rain forest, 24 temperate forest and desert Figure Proportions of functional groups in subtropical bushland, 25 montane forest and tropical grassland Figure Relationships between proportions of functional groups and age 26 of forests Figure Relationships between proportions of functional groups and age 27 of forests Figure Correlations of proportions of Opportunist with Generalized 28 Myrmicinae, Cryptic Species and Specialized Predator ants Figure Correlations of Cryptic Species with Tropical Climate Specialist, 29 Generalized Myrmicinae and Specialized Predator ants Figure Correlations between proportions of functional groups with one 30 another Figure Correlation between proportions of Tropical Climate Specialist 31 and Specialized Predator ants Figure Relationship between proportions of Cold Climate Specialist and 31 age of temperate forests Figure 10 Sigmoid curves relating the proportions of functional groups and 32 the age of tropical rain forests Figure 11 Sketch map of Singapore main island 45 v Figure 12 Study sites and transects in the Central Catchment Nature 53 Reserve Figure 13 Correlation between proportions of functional groups and the 58 ranked age of Forest Type Figure 14 Correlation between proportions of functional groups and the 59 ranked age of Forest Type Figure 15 Correlations of proportions of Cryptic Species, Generalized 60 Myrmicinae and Specialized Predator groups with Opportunist group Figure 16 Correlations of proportions of Cryptic Species, Generalized 61 Myrmicinae and Specialized Predator groups with Tropical Climate Specialist group Figure 17 NMS plots of species and site scores 64 Figure 18 Estimated mean age of different Forest Types 65 vi CHAPTER GENERAL INTRODUCTION Succession Ecology Ecosystems are not static entities Conditions within habitats vary widely and one group of these variations is ecological succession cycles For example, large trees in primary rain forests often collapse, leaving gaps in the canopy and creating patches where secondary succession takes place (Whitmore, 1998) The gaps slowly return to the primary condition after many years, completing the cycle (Whitmore, 1998) There are similar cycles in other areas such as aquatic habitats (Wetzel, 1995) The general trend seen in ecological succession is that of particular community structure and environmental conditions replacing another over time There usually is a set of environmental conditions that allow a certain community to thrive As this community ages, the conditions change and another set of community more suited to the new conditions gradually take over (McCook, 1994) Furthermore, it is not just the interactions between environment and community that play important roles Interactions between different species are also important in determining the path of succession (Farrell, 1991) In addition, one of the most important determinants of whether an ecosystem recovers naturally is the disturbance caused by humans The over-riding significance of the ecological footprints created by anthropogenic disturbance is clearly demonstrated in large-scale landscape changes brought about through the advancement and development of human civilization (DeFries et al, 2005) The natural cycles are either altered or replaced by artificial cycles For example, cultivated landscapes that were subsequently abandoned often not have the conditions that are necessary for growth into its original uncultivated state (Roth, 1999) The natural resources of this planet are limited but yet are being exploited at a high rate (DeFries et al, 2005) There is an urgent need to restore exploited ecosystems and bring it back from the brink of collapse This is important because ecosystems provide numerous services (Bennett et al, 2005) Having a good understanding of succession ecology will enable stakeholders of the environment to effectively rehabilitate and conserve natural resources so that sustainable use is achieved for the long term (Leitao and Ahern, 2002) Many ecologists try to grasp the complexity of succession by describing its individual components and making useful extrapolations in order to understand general trends (Underwood, 1997) Another challenge that ecologists deal with is to come up with reliable models of prediction from their observations Predictive models in ecology are useful in that it allows researchers to work on a general level with fewer variables to estimate the responses of ecosystems (Brook et al, 2000) Such models could serve as an alert mechanism for possible impending ecological crisis, much like climate change and global warming models (Viner et al, 1995) One of the components of succession is the animal community The fauna of a habitat changes with the succession cycle (Done, 1992) Therefore, ecologists often study changes in populations and communities of animals that affect and are themselves affected by succession For example, the community structure, species richness and abundance are different at different stages of succession (Done, 1992) Ants are considered a useful group of organisms for the study of ecosystem succession This is because of their relatively wide ecological range (Holldobler and Wilson, 1990) and the different responses of different species to changes in the habitats (Andersen, 1995) Aspects of ant biology and their significance in ecosystems will be discussed shortly Biology of The Ants “Ants are everywhere,” Holldobler and Wilson (1990) once commented It is not surprising that such a statement was made Ants are insects, which constitute over 75% of all estimated animal species on this planet (Ruppert and Barnes, 1994) There are more than 9,000 described species of ants in nearly 300 genera, forming the entire family Formicidae, within the order Hymenoptera (Bolton, 1994) There are three anatomical features found in the ant that set it apart from other insects First, ants have narrow proximal segments of the abdomen joining the thorax These narrow segments are known collectively as the petiole (Bolton, 1994) Second, the mandibles of an ant 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Harvard University Press, Cambridge, Massachusetts Wong, P P 1968 Some aspects of the surface configuration of Singapore island M.A 93 thesis, National University of Singapore Wong, Y K., Chew, P T and Ali, I 1994 The tree communities of the Central Catchment Nature Reserve, Singapore Gardens’ Bulletin Singapore 46: 37-78 Yamane, S., Itino, T & Nona, A R 1996 Ground ant fauna in a Bornean dipterocarp forest The Raffles bulletin of zoology 44: 253-262 Zar, J H 1999 Biostatistical analysis (4th edition) Prentice Hall, New Jersey 94 APPENDIX A List of ant genera with functional group classification (from Brown, 2000) CCS = Cold Climate Specialist, CS = Cryptic Species, DD = Dominant Dolichoderinae, GM = Generalized Myrmicinae, HCS = Hot Climate Specialist, OP = Opportunist, SC = Subdominant Camponotini, SP = Specialized Predator, TCS = Tropical Climate Specialist Genus Acanthognathus Acanthomyops Acanthomyrmex Acanthoponera Acanthostichus Acromyrmex Acropyga Adelomyrmex Adlerzia Aenictogiton Aenictus Afroxyridris Allomerus Amblyopone Anillomyrma Anisopheidole Ankylomyrma Anochetus Anomalomyrma Anonychomyrma Anoplolepis Aphaenogaster Aphomomyrmex Apomyrma Apterostigma Asphinctanillodes Asphinctopone Atopomyrmex Atta Azteca Baracidris Basiceros Belonopelta Blepheridatta Functional group SP CS TCS TCS CS TCS CS TCS CS TCS TCS CS CS CS CS CS TCS SP CS DD CS OP TCS CS TCS CS CS TCS TCS DD CS CS CS TCS 95 Bondriotia Bothriomyrmex Bracymyrmex Calomyrmex Calyptomyrmex Camponotus Cardiocondyla Carebara Carebarella Cataglyphis Cataulacus Centromyrmex Cephalotes Cerapachys Chalepoxenus Cheliomyrmex Chimaeridris Cladomyrma Colobostruma Concotio Creightonidris Crematogaster Cryptopone Cylindromyrmex Cyphoidris Cyphomyrmex Dacetinops Daceton Dendromyrmex Diacamma Dicroaspis Dilobocondyla Dinoponera Discothyrea Doleromyrma Dolichoderus Doronomyrmex Dorylus Dorymyrmex Dysedrognathus Echinopla Eciton Ectatomma Emerypone Epelysidris Epimyrma Epitrius Epopostruma CS CCS TCS CS CS SC OP CS CS HCS TCS CS TCS CS CCS TCS TCS TCS SP CS CS GM CS SP TCS TCS CS SP SC OP CS TCS SP CS OP TCS CCS TCS OP CS SC TCS OP CS TCS CCS CS SP 96 Eucryptocerus Euprenolepis Eurhopalothrix Forelius Forelophilus Formica Formicoxenus Froggetella Gesomyrmex Gigantiops Glamyromyrmex Gnamptogenys Goniomma Gymnomyrmex Harpagoxenus Harpegnathos Heteroponera Huberia Hylomyrma Hypoponera Indomyrma Ireneopone Iridomyrmex Ishakidris Kartidris Kyidris Labidus Lachnomyrmex Lasiophanes Lasius Leptanilla Leptanilloides Leptogenys Leptomyrmex Leptothorax Linepithema Liometopum Liomyrmex Lophomyrmex Lordomyrma Lordomyrmex Loweriella Machomyrma Manica Mayriella Megalomyrmex Melophorus Meranoplus TCS TCS CS DD SC CCS CCS DD TCS TCS CS TCS HCS CS CCS SP CCS CCS TCS CS CS TCS DD CS TCS CS TCS TCS CCS CCS CS CS SP TCS TCS DD DD TCS TCS TCS TCS TCS CS CCS CS TCS HCS HCS 97 Mesostruma Messor Metapone Metapone Microdaceton Monomorium Mycetarotes Mycetophylax Mycetosoritis Mycocepurus Myopias Myopopone Myrcidris Myrmecia Myrmecina Myrmecocystus Myrmecorhynchus Myrmelachista Myrmica Myrmicaria Myrmicaria Myrmicocrypta Myrmoteras Mystrium Neivamyrmex Neostruma Nomamyrmex Noonilla Nothidris Nothomyrmecia Notoncus Notostigma Ochetellus Ochetomyrmex Octostruma Ocymyrmex Odontomachus Odontoponera Oecophylla Oligomyrmex Onychomyrmex Opisthopsis Orectognathus Overbeckia Oxyepoecus Oxyopomyrmex Pachycondyla Paedalgus SP HCS TCS TCS CS GM TCS TCS TCS TCS CS CS TCS SP TCS HCS CCS CS OP TCS TCS TCS SP CS TCS CS TCS CS CCS SP CCS SC OP TCS CS HCS OP SP TCS CS TCS SC SP SC TCS HCS SP CS 98 Papyrius Paratopula Paratrechina Pentastruma Perissiomyrmex Peronomyrmex Petalomyrmex Phasmomyrmex Phaulomyrma Pheidole Pheidologeton Philidris Phrynoponera Plagiolepis Platythyrea Plectroctena Podomyrma Poecilimyrma Pogonomyrmex Polyergus Polyrhachis Ponera Prenolepis Prionopelta Pristomyrmex Proatta Probolomyrmex Proceratium Procryptocerus Prolasius Protalaridris Protanilla Protomognathus Psalidomyrmex Pseudaphomomyrmex Pseudoatta Pseudolasius Pseudomyrmex Pseudonotoncus Quadristruma Recurvidris Rhopalothrix Rhytidoponera Rogeria Romblonella Rotastruma Santschiella Scyphodon DD TCS OP CS TCS TCS TCS SC CS GM CS DD SP CS SP SP TCS TCS HCS SP SC CS CCS CS TCS TCS CS CS TCS CCS CS CS CCS CS TCS TCS TCS TCS CCS CS CS CS OP TCS TCS TCS TCS CS 99 Sericomyrmex Serrastruma Simopelta Simopone Smithstruma Solenopsis Sphinctomyrmex Stegomyrmex Stenamma Stereomyrmex Stigmacros Streblognathus Strumigenys Talaridris Tapinoma Tatudris Technomyrmex Terataner Teratomyrmex Tetramorium Tetraponera Tetraponera Thaumatomyrmex Tingimyrmex Trachymyrmex Tranopelta Trichoscapa Tricytarus Turneria Typhlomyrmex Vollenhovia Vombisidris Wasmannia Xenomyrmex Yavnella Zacryotocerus TCS CS CS SP CS CS CS CS CCS TCS CCS SP CS CS OP CS OP TCS TCS OP TCS TCS SP CS TCS CS CS TCS TCS CS TCS TCS TCS TCS CS TCS 100 APPENDIX B List of the number of ant individuals collected in the four Forest Types in Singapore CCS = Cold Climate Specialist, CS = Cryptic Species, DD = Dominant Dolichoderinae, GM = Generalized Myrmicinae, HCS = Hot Climate Specialist, OP = Opportunist, SC = Subdominant Camponotini, SP = Specialized Predator, TCS = Tropical Climate Specialist Species Acanthomyrmex sp Acropyga sp Aenictus sp Anochetus sp Anoplolepis sp Aphaenogaster sp Camponotus gigas Camponotus sp Camponotus sp Camponotus sp Cardiocondyla sp Cerapachys sp Crematogaster sp Crematogaster sp Diacamma sp Dolichoderus sp Dolichoderus sp Dolichoderus sp Euprenolepis sp Gnamptogenys sp Hypoponera sp Hypoponera sp Iridomyrmex sp Leptogenys sp Leptogenys sp Leptogenys sp Lophomyrmex sp Meranoplus sp Metapone sp Monomorium sp Myrmicaria sp Ochetellus sp Odontomachus sp 1 0 2 0 2 3 0 1 29 Forest Type 1 3 11 13 12 14 5 14 12 12 10 29 12 11 9 Functional Group 58 12 11 1 1 10 18 17 1 3 TCS CS TCS SP CS OP SC SC SC SC OP CS GM GM OP TCS TCS TCS TCS TCS CS CS DD SP SP SP TCS HCS TCS GM TCS OP OP 101 Odontomachus sp Odontoponera sp Odontoponera sp Opisthopsis sp Pachycondyla sp Paratopula sp Paratrechina sp Pheidole sp Pheidole sp Pheidole sp Pheidole sp Pheidole sp Pheidole sp Pheidole sp Pheidole sp Pheidole sp 10 Pheidole sp 11 Pheidole sp 12 Pheidole sp 13 Pheidole sp 14 Pheidole sp 15 Pheidole sp 16 Pheidole sp 17 Pheidologeton sp Pheidologeton sp Philidris sp Poecilimyrma sp Polyrhachis sp Prenolepis sp Proatta sp Pseudolasius sp Strumigenys sp Technomyrmex sp Tetramorium sp Tetramorium sp Tetramorium sp Tetramorium sp Tetraponera sp Vollenhovia sp Wasmannia sp Total 1 0 1 0 0 0 0 0 2 0 105 0 12 3 4 20 10 13 12 10 1 11 14 19 441 35 10 1 11 35 2 23 10 51 39 19 16 15 1 1 462 12 20 22 3 34 15 13 6 42 10 27 13 1 1 0 464 OP SP SP SC SP TCS OP GM GM GM GM GM GM GM GM GM GM GM GM GM GM GM GM CS CS DD TCS SC CCS TCS TCS CS OP OP OP OP OP TCS TCS TCS 102