Species Diversity of Terrestrial Earthworms in Different
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SPECIES DIVERSITY OF TERRESTRIAL EARTHWORMS IN KHAO YAI NATIONAL PARK Prasuk Kosavititkul A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Environmental Biology Suranaree University of Technology Academic Year 2005 ISBN 974-533-516-9 ความหลากหลายของชนิดไสเดือนดินในเขตอุทยานแหงชาติเขาใหญ นายประสุข โฆษวิฑิตกุล วิทยานิพนธน เปี้ นสวนหนงของการศึ่ ึกษาตามหลักสูตรปริญญาวิทยาศาสตรดุษฎีบณฑั ิต สาขาวิชาชีววทยาสิ ิ่งแวดลอม มหาวิทยาลัยเทคโนโลยีสุรนารี ปการศึกษา 2548 ISBN 974-533-516-9 ACKNOWLEDGEMENTS I am most grateful to thank Asst. Prof. Dr. Panee Wannitikul, my advisor, for her generous help, encouragement and guidance throughout this thesis from the beginning. Her criticism, improvement, and proper of manuscript have made this thesis in correct form. I sincerely thank to Assoc. Prof. Dr. Korakod Indrapichate, Assoc. Prof. Dr. Somsak Panha, Dr. Nathawut Thanee and Dr. Pongthep Suwanwaree for their valuable advice and guidance in this thesis. I am sincerely grateful to Assoc. Prof. Dr. Sam James, Natural History Museum and Biodiversity Research Center, Kansas University for his helpful and kind assistance in confirming and identifying earthworms. Thanks also to the Center for Scientific and Technological Equipment, Suranaree University of Technology for the laboratory facilities and scientific instruments. Special thank is due to the Khao Yai National Park for permitting me to work in the park. I would like to express my special thank to Naresuan University for the scholarship supporting my study. Special gratitude is expressed to my parents, my sisters, my family, my friends, my seniors, my juniors, and other people who give me a supported power whenever I lose my own power. Prasuk Kosavititkul CONTENTS Page ABSTRACT IN THAI…………………………………………………………… I ABSTRACT IN ENGLISH……………………………………………………… II ACKNOWLEDGEMENTS……………………………………………………... III CONTENTS……………………………………………………………………… IV LIST OF TABLES………………………………………………………………. VII LIST OF FIGURES……………………………………………………………... VIII CHAPTER I INTRODUCTION 1.1 The importance of problems………………………………………… 1 1.2 The objectives of study……………………………………………… 2 1.3 The scope and limitations of study………………………………….. 2 1.4 Key words ………………………………………………………….... 3 1.5 Details of the study area……………………………………………… 3 II LITERATURE REVIEW 2.1 External characteristics……………………………………………… 7 2.2 Internal anatomy and function………………………………………. 15 2.3 Earthworm diversity and geographical distribution………………..... 23 2.4 Earthworm ecology………………………………………………….. 30 2.5 The influence of environmental factors on earthworms…………….. 35 V CONTENTS (Continued) Page III RESEARCH METHOD 3.1 Study site description………………………………………………. 43 3.2 Climate……………………………………………………………... 45 3.3 Earthworm sampling and identification……………………………. 45 3.4 Litter sampling……………………………………………………... 48 3.5 Soil sampling and analysis…………………………………………. 48 3.6 Data analysis………………………………………………………… 49 IV RESULTS AND DISCUSSION 4.1 Environmental factors……………………………………………… 52 4.2 Earthworm identification…………………………………………… 58 4.3 Earthworm community and distribution……………………………. 101 4.4 Stepwise multiple regression analysis of earthworm community…………………………………………………………. 120 V CONCLUSION 5.1 Conclusion………………………………………………………….. 124 5.2 Recommendation…………………………………………………… 125 REFERENCES………………………………………………………………… 126 APPENDICES APPENDIX A Definitions…………………………………………….. 150 APPENDIX B Number and species of earthworms in five forest types…………………………………………….. 158 VI CONTENTS (Continued) Page APPENDIX C Environmental factors in five forest types……………. 174 CURRICULUM VITAE………………………………………… LIST OF TABLES Table Page 4.1 Mean and standard error (±SE) of climate factors, litter and soil properties of five forest types……………………………………… 53 4.2 Occurrence of families and species of earthworms collected in Khao Yai National Park…………………………………………….. 59 4.3 Occurrence of species and numbers of earthworms collected in five forest types…………………………………………………….. 102 4.4 The abundance of earthworm species in July-December 2002 and June-December 2003 in sampling areas in Khao Yai National Park… 109 4.5 The density of earthworm species in MD, DE, ME, HE and GL……. 112 4.6 Species diversity index and evenness index of earthworms in five forest types………………………………………………………. 113 4.7 Similarity index of earthworm community of Khao Yai National Park…………………………………………………………………... 114 4.8 The Pearson and Kendall correlation with ordination axis………… . 118 LIST OF FIGURES Figure Page 2.1 Ventral view of external anatomy of earthworms…………………… 8 2.2 Various forms of prostomium……………………………………….. 8 2.3 Arrangement of setae in Oligochaeta………………………………. .. 10 2.4 Penial setae of some Australian Megascolecidae……………………. 10 2.5 Cocoon formation and arrangement of male and prostatic pores in the Megascolecidae………………………………………………….. 12 2.6 An example of a megascolecine genital field……………………….. 14 2.7 Digestive, circulatory and reproductive systems of earthworms……. 17 2.8 Calciferous glands and their vascularization in some Australian Megascolecidae……………………………………………………… 18 2.9 Vascular system. Dorsal blood vessel and connections in the oesophageal region of Megascolecidae exemplified by Didymogaster sylvaticus ……………………………………………. 19 2.10 The genital organs of a megascolecine…………………………........ 23 3.1 The map of Khao Yai Nation Park shows 5 different sampling areas.. 44 3.2 Sampling grid design used to sample the earthworm community within each quadrat…………………………………………………… 47 IX LIST OF FIGURES (Continued) Figure Page 4.1 Average of rainfall (mm) from 2002 to 2003 at Pak Chong Meteorological Station, Nakhon Ratchasima Province……….…………. 54 4.2 Photographs and line drawings of Pontoscolex corethrurus…………….. 63 4.3 Photographs and line drawings of Perionyx excavatus………………….. 67 4.4 Photographs and line drawings of Amynthas alexandri…………………. 70 4.5 Photographs and line drawings of Amynthas fucosus…………………… 73 4.6 Photographs and line drawings of Amynthas longicauliculatus………… 75 4.7 Photographs and line drawings of Amynthas sp. 1………………………. 77 4.8 Photographs and line drawings of Amynthas sp. 2………………………. 80 4.9 Photographs and line drawings of Amynthas sp. 3………………………. 82 4.10 Photographs and line drawings of Metaphire peguana…………………. 86 4.11 Photographs and line drawings of Metaphire houlleti………………….. 89 4.12 Photographs and line drawings of Metaphire sp.1………………………. 91 4.13 Photographs and line drawings of Pithemera bicincta………………….. 94 4.14 Photographs and line drawings of Drawida beddardi…………………... 99 4.15 The abundance of earthworm composition, juvenile, subadult and adult earthworms , and cocoon of P. corethrurus, in July-December 2002 and June-December 2003 in sampling areas in Khao Yai National Park……………………………………………………………………. 106 X LIST OF FIGURES (Continued) Figure Page 4.16 The abundance of earthworms in five forest types in Khao Yai National Park…………………………………………………………. 111 4.17 Dendrogram for hierarchical clustering of earthworms in five forest types…………………………………………………………… 115 4.18 The joint plot diagram showing the relationship between a set of ecological factors and earthworm abundance……………………….. 116 CHAPTER I INTRODUCTION 1.1 The importance of problems Earthworms are classified in phylum Annelida, class Oligochaeta. They are presumably the most ancient soil dwellers and were found earlier as in sediments of the Pre-Cambrian and Ordovician (Miller and Harley, 1999). The most recent data account for 3627 species described worldwide (Reynolds, 1994). Charles Darwin was one of the first scientists to give credence to the conventional wisdom from earlier civilizations about the importance of earthworms to soil fertility. Recently there has been resurgence in interest in earthworms, driven by environmental and economic concerns, particularly the need to understand and utilize their functions in sustainable agriculture, and to exploit their potential for recycling of organic wastes (Lee, 1985; Sims and Garard, 1985). Earthworms are important in the breakdown of organic matter to inorganic matter. The action of the digestive fluids and increased microbial activity in the casts tends to solubilize inorganic plant nutrient elements. The extensive burrowing of the earthworm improves soil aeration and increases water penetration into soil. Upon further microbial decomposition of the partially digested residues, plant nutrient elements are released (Edwards, 1998). Thailand is located in the tropical region which encompasses diverse kinds of natural ecosystems. These natural habitats are homes to some of the world’s richest 2 and unique plants and animals, resulting in a high diversity of earthworms. However, the studies in biology and ecology dealing with earthworms in Thailand have been lagging behind due to the limited knowledge of both their morphology and taxonomy. This investigation will provide information of earthworms on species compositions, quantities, abundance, earthworm diversity, seasonal variation and some ecological factors effecting earthworm compositions. Information from this study will also increase knowledge and understanding of ecosystem changes, and be making its primary contributions to the work of people whose interests line in ecology, agriculture, and other fields in need of a coherent classification of earthworm species. 1.2 The objectives of study 1.2.1 To investigate species diversity of terrestrial earthworms in different vegetation types (mixed deciduous forest, dry evergreen forest, moist evergreen forest, hill evergreen forest and grassland) in Khao Yai National Park.