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TAXONOMY and ECOLOGY of NEMATODES OFAGRICULTURAL FIELDS and WASTELANDS Doctor of Philosophy ZOOLOGY GAURAV KUMAR SINGH 2011

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TAXONOMY and ECOLOGY of NEMATODES OFAGRICULTURAL FIELDS and WASTELANDS Doctor of Philosophy ZOOLOGY GAURAV KUMAR SINGH 2011 TAXONOMY AND ECOLOGY OF NEMATODES OFAGRICULTURAL FIELDS AND WASTELANDS THESIS SUBMITTED FOR THE AWARD OF THE DEGREE OF Doctor of Philosophy IN ZOOLOGY BY GAURAV KUMAR SINGH DEPARTMENT OF ZOOLOGY ALIGARH MUSLIM UNIVERSITY ALIGARH (INDIA) 2011 Wxw|vtàxw gÉ `ç UxÄÉäxw ctÜxÇàá IRFAN AHMAD Ph.D. DEPARTMENT OF ZOOLOGY Professor Aligarh Muslim University, Aligarh – 202002, INDIA [email protected] Date: …………………. Certificate This is to certify that the research work presented in the thesis entitled, “Taxonomy and Ecology of Nematodes of Agricultural Fields and Wastelands” by Mr. Gaurav Kumar Singh is original and was carried out under my supervision. I have permitted Mr. Singh to submit it to the Aligarh Muslim University, Aligarh in fulfillment of the requirements for the degree of Doctor of Philosophy in Zoology. IRFAN AHMAD (Supervisor) Acknowledgements I bow in deep reverence to GOD the Almighty, who blessed me with an innumerable favour of academic work. It has been a great opportunity for me to work under the able guidance of Prof. Irfan Ahmad, Chairman, Department of Zoology, Aligarh Muslim University, Aligarh. I express my sincere gratitude for his ever-lasting important advices, valuable suggestions, stimulating discussions, constructive criticism, sense of perfection and precision which enabled me to complete this work. Inspite of his tight departmental commitments he kept his door open wide for me. His affectionate instinct and constant encouragement were always a boon to me. Any error if still remains is entirely my own. I am thankful to the Chairmen (past and present), Department of Zoology, Aligarh Muslim University, Aligarh for providing necessary laboratory facilities for the work. I feel more than obliged to all my respected teachers Prof. M. Shamim Jairajpuri (INSA Senior Scientist), Prof. Wasim Ahmad, Prof. Qudsia Tahseen and Prof. Mahalaqa Choudhary for their valuable advices and kind suggestions. I wish to acknowledge my seniors and lab colleagues, Dr. Noorus Sabah, Dr. Md. Mahamood, Dr. Ali Asghar Shah, Mr. Puneet Kumar, Ms. Gazala Yousuf & Ms. Nadia Sufyan. I am also thankful to my senior Dr. Md. Baniyamuddin and colleagues Dr. Shikha Ahlawat, Ms. Uzma Tauheed, Mrs. Tabinda Nusrat , Ms. Malka Mustaqim and Ms. Sumaiya Ahad for their constant inspirations and support. Special thanks are due to my friend and colleague Vijay Vikram Singh for his unconditional and constant help during compilation of this work. Thanks are also due to my friends Imran and Vimal for their constant encouragement during the course of present work. Last but not the least, I would like to thank my wife for making me believe that I can do it and my little angel for giving me cheers of my life. My brothers Mr. Umesh Singh and Mr. Saurabh and my bhabhi Mrs. Meenakshi deserve special thanks for creating ideal milieu at home which helped me to complete the present work. The financial assistance from the Ministry of Environment and Forest, Government of India, New Delhi is also thankfully acknowledged. Gaurav Kumar Singh CONTENTS PART – A Page INTRODUCTION 1 HISTORICAL BACKGROUND 12 MATERIALS AND METHODS 21 SYSTEMATICS 24 ORDER RHABDITIDA 24 SUBORDER CEPHALOBINA 24 SUPERFAMILY CEPHALOBOIDEA 25 FAMILY CEPHALOBIDAE 26 SUBFAMILY CEPHALOBINAE 26 Genus Pseudacrobeles 27 Pseudacrobeles ventricauda sp. n. 28 Pseudacrobeles mucronatus sp. n. 34 SUBFAMILY ACROBELINAE 41 Genus Acrobeles 42 Acrobeles mariannae 42 Genus Acrobeloides 47 Acrobeloides glandulatus sp. n. 47 Genus Cervidellus 54 Cervidellus neoalutus sp. n. 54 Cervidellus minutus sp. n. 60 Genus Chiloplacus 66 Chiloplacus aligarhensis sp. n. 66 Genus Nothacrobeles 73 Nothacrobeles punctatus sp.n. 73 Genus Stegellata 80 Stegellata ophioglossa 80 Genus Zeldia 85 Zeldia tridentata 85 SUPERFAMILY PANAGROLAIMOIDEA 90 FAMILY PANAGROLAIMIDAE 90 SUBFAMILY TRICEPHALOBINAE 91 Genus Tricephalobus 91 Tricephalobus quadripapilli sp. n. 92 FAMILY BREVIBUCCIDAE 98 Subfamily Brevibuccinae 98 Genus Brevibucca 98 Brevibucca postamphidia sp. n 99 Genus Plectonchus 105 Plectonchus coptaxii sp. n. 105 SUMMARY 112 PART – B INTRODUCTION 115 MATERIALS AND METHODS 125 RESULTS 131 DISCUSSION 153 REFERENCES 162 Part – A Taxonomy Introduction The nematodes are a successful group of invertebrates placed at a low level in the taxonomic hierarchy of the animal kingdom. They are the most diverse phylum, and one of the most diverse of all animals. They have successfully adapted to nearly every ecological habitat from marine to fresh water, from the Polar regions to the tropics, as well as the highest to the lowest of elevations. They are ubiquitous in freshwater, marine, and terrestrial environments, where they often outnumber other animals in both individual and species counts, and are found in the locations as diverse as Antarctica and oceanic trenches. Some of them also can withstand complete dryness on the surface of rocks. Their size too is extremely variable ranging from less than 100 µm (Greefiella minutum) to greater than 8 metres (Placentonema gigantissima). The nematodes are the planets most abundant metazoa; of every five animals, four are nematodes (Platt, 1994; Bongers & Ferris, 1999). They are particularly abundant in marine, freshwater, and soil habitats. A square yard of woodland or agricultural habitat may contain several million nematodes. The existence of nematodes living in water, soil or in parasitizing the plants remained largely unknown perhaps, because of their exceedingly small size, absence of any colouration, mostly underground habitat and the difficulties encountered in their isolation. The phylum Nematoda is characterized by high species diversity. It has been estimated that the total number of described and undescribed species might be ranged from 0.1 to 100 million (May, 1988; Hammond, 1992; Lambshead,1993; Coomans, 2000). The nematodes are not only numerically 1 abundant, but they are also very diverse in terms of species. Usually species richness at a single site is high with an average of 20-60 species per soil sample. In soil, the nematodes dominate in number as well as species over all other soil inhabiting animals collectively and have occupied all possible habitats representing a very wide range of biological diversity. Soil nematode communities have the potential to provide insights into many soil processes and functions as most nematodes are active in soil throughout the year (Ritz & Trudgill, 1999). Nematodes can be used as bioindicators of soil health because they are ubiquitous and have diverse feeding behaviours and life strategies (Bongers & Bongers, 1998; Neher, 2001). They occupy several trophic grades and a central position in the soil food web and play significant roles in biological processes such as nitrogen cycling and plant growth patterns (Neher, 2001). Soil nematodes stabilize soil ecosystems, promote substance cycling and energy flow (Ingham et al., 1985). The ecological classification of terrestrial nematodes have usually been based on feeding biology (trophic functions) and on life strategies; colonizers versus persisters (Bongers, 1990). Yeates et al. (1993) classified the terrestrial nematodes into eight trophic groups viz., plant feeding, hyphal feeding, substrate ingestion, predation on animals, bacterial feeding, unicellular eukaryote feeding, dispersal/infective stage of parasites and omnivorous. Free-living nematodes promote decomposition of soil organic matter, mineralization of plant nutrients and nutrient cycling, amend soil physico-chemical property and improve soil fertility (Ferris et al., 2004). Some free living nematodes suppress bacterial, fungal and nematode diseases (Khan & Kim, 2007). 2 All terrestrial ecosystems consist of aboveground and belowground components that interact to influence community- and ecosystem-level processes. Several recent studies have indicated that biotic interactions in soil can regulate the structure and functioning of the above ground communities (Wardle et al., 2004). Soil nematode communities can provide unique insights into many aspects of soil processes; they can offer a holistic measure of the biotic and functional status of soils (Ritz & Trudgill, 1999). They are good environmental indicators because of their strong relationships with land management (Todd, 1996; Neher and Campbell, 1994; Liang et al., 1999; Fiscus and Neher, 2002) and aboveground vegetation (Ingham et al., 1985; Bongers and Bongers, 1998; Bongers and Ferris, 1999; Yeates, 1999). Measurement of meiofaunal diversity and abundance is an important but time consuming process. Morphological identification of individual organisms to named species is often not technically possible due to sheer abundance, small size, and lack of expert knowledge of the groups encountered. This is especially true of nematodes, whose diversity in soils and sediments remains essentially unknown. Surveys of benthic sediments suggest that the total species number for marine nematodes may exceed 1 million (Lambshead 1993; Lambshead 2001), with only a few thousand described in the scientific literature (Malakhov 1994; De Ley & Blaxter 2002). In terrestrial systems, nematode diversity appears to be under-reported (Lawton et al., 1998), with, for example, only about 200 species of soil nematodes being described from the British Isles (Boag & Yeates 1998). The maximum number of nematode taxa described from a single soil site is 228 from a prairie in Kansas, USA (Orr & Dickerson 1966; Boag & Yeates 1998). 3 Nematodes are arguably the most numerous metazoans in the soil and aquatic
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