STUDIES on the TAXONOMY and BIOLOGY of SOME DIPLOGASTRID SPECIES from INDIA Tdoctor of Pbili)!
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STUDIES ON THE TAXONOMY AND BIOLOGY OF SOME DIPLOGASTRID SPECIES FROM INDIA THESIS SUBMITTED FOR THE AWARD OF THE DEGREE OF TDoctor of Pbili)!gopbp IN ZOOLOGY BY GAZALA YOUSUF DEPARTMENT OF ZOOLOGY ALIGARH MUSLIM UNIVERSITY ALIGARH -202002, INDIA .- C1S 2013 .c~5 iad f rh, BOO 13 ,6V 2014 IIIIIIII III IIIII illill IIIII lillil!l T9005 , IRFAN AHMAD Ph.D. DEPARTMENT OF ZOOLOGY Professor Aligarh Muslim University, Aligarh — 202002, INDIA ahmadirfi c,gmaiI.com Date: ...................... Certificate This is to certify that the research work embodied in the thesis entitled, "Studies on the Taxonomy and Biology of some Diplogastrid species from India" by Ms. Gazala Yousuf is original and was carried out under my supervision. I have permitted Ms. Gazala to submit it to the Aligarh Muslim University, Aligarh in fulfillment of the requirements for the degree of Doctor of Philosophy in Zoology. IRFAN ARMAD (Supervisor) )lcknow(ecfgements 6vu to deep re(ereucc to the Altrtig(ty A(Kih, the most 6eneficia(and mercifu( ,whose benign blessings led me to complete this wcvorl,and produce it in its present fonn. From the core of my heart, I express my deep sense of gratitude to my supertasor Prof Irfan Ahmad, Chairman, •Department of ."oolojy, aI' -Dean, 'Facu(ty of Life Sciences, /1(efjarh :tfus(unt Z'nicersity, A(igarfr for his eter-fasting important ad.ices, valuable suggestions, stimulating discussions, constructice criticis►n, sense of perfection and precision tchic( enabled me to complete this tror1,. Inspite of his tight departmental commitments he (dept his door wide open for ire. I(is affectionate instinct and constant encouragement were always a boon to me. It has been an honour to to orry with him. I also tharva ,Prt!..11. S(antun ,jairajpuri r1'\;5 ,Senior, Scientist I, Prof •Rasi►n )tlztmad Prof Qudsia lahsee►t and Pro,;%%faha(aga (Youdhary for their valuable advices and kind suggestions. Helping hands extended by my seniors and (a6 colleagues, Dr. Moorus ,Sa6a(e, Or Md ufahamood, ,Dr. Gauraz Singh, ;Mr. (Puneet Kumar e Wrs..A'adia Sufyart are highly acknoufedged here with thanIs. I am also t(tanGfu(to my seniors Dr. ifd Baniyamuddirt, Dr. S(ikha Al fa-wat, Dr. 'Vyay Yikram Singh, Sots. Z'zma Tauheed, wr. i4d )lsif, Efs. Yaf&a 'Wustagirn, .Ms. Sumaya)4had and 11s. Zarrn Imran for their pleasant company, help and support. Special thanks are due to 'Late) _Mrs. '1"abinda Ivusrat for her moral support, suggestions and above all good company. She a as more than just a friend and the coif she (eft in my heart can serer be fiffed: I tuou(d file to proride a token off thanks to my friends Azf a, Seema, ifuma, Sadaf, Xqzia, %al wish and'7' a.ia, a (to though not a part of this research i ork. but supported me to an a ytrerne extent which I can't forget. ?flanks are also due to my class-mates ;Ifafino, 7alak, 'Farha, 'Yusra and Waseem for their help and support. My words of thanks will remain incomplete if I forget to mention the names of my friends 'Fatima, 7fumaira andSumaira, who were oath me during the ups and downs of my research, gave me their precious time, patient hearing and kindness during the days I really found difficult to carry out my workand sorted out my pro6iems. Last but not the (east, I wouldG&e to thank my Parents, my elder Brothers Mr. 'Khal dlousuf a ;Nr. ;NubashIr ;Yousu f my Sister-in-law ;%1rs.:Naroof Aaiir for their unconditional support, loco and never ending care. I am equally thankfu( to my little niece Tahura BtnteN,ha(id forgiBang me cheers of my life. Gaala Yousuf CONTENTS Page Introduction I IIistorical Background 7 Taxonomic revic« of free-living nematodes 7 Re% ie%% on biology:) of free-living nematodes 30 Materials and Methods 42 Systematics 46 Order Rhanditida 46 Suborder 1)iplogastrina 46 Superfamii> Diplogastroidea 47 amih Diplogasteroididae 48 SuhfamiI 1)iplogasteroidinae -48 Genus (;offurtiu 48 Go fiirtiu bcrlchus sp.n. 49 Subfamily Rhabditolaiminae 55 Genus Rhandliticloicic.s Rhuhditidoi(l s hapillutu.t 5 5 Family Diplogastridae 61 Subfamily Diplogastrinae 61 Genus Aerosviehus 6I Acrostichuc nudicupitutus 62 Genus Butlerius 68 Butlerius micuns 68 Genus Iiplogu.vtrellus 74 I Diplogasirellus curvatus sp. n. 74 Diplogastrellus neograciloides sp. n. 80 lli1,lr,ga.c•trdlIus ka.shmerieasis sp. n. 86 Diploc astrellus loktakai sp. n. 92 Diploga.strellus gracilis 98 Genus I'serrckuvusticlt,c.s gen. n. 103 I'.%rtnk,c'ruslirhuo ,uu.sal sp. n. I04 Subfamily Ihmaniellinac 111 Genus Dernaniellu 1 l 1 Demaniella keirakensis l 1 1 Family Neodiplogastridae 117 Subfamily Neodiplogastrinae 117 Genus Fictor 118 bictor suptilts sp. n. 118 Fictor 1aj2giruac!u sp. n. 125 Genus Koerneria 131 Koerl1e!ia/llicaudala 131 Genus 1!fononc1roirlc.c 137 Afononchoick's (enuispu•u!a sp. n. 137 .%fononchoide.c latiguhernacula sp. n. 144 Afononc/,oicic'.s .sn•uitu.c 1411 Genus ()igolaimella 156 Oigolaimella breviguhernacula sp. n. 156 Oigolaiacl!a longic•auda 162 Genus I'ris1ion{lu.c 167 l'risiionc/ics stercnrari.s sp. n. 167 I mil%"I\IL~pharyngidae 174 Genus 7}ioplttol•itr 17.1 71 lop/tarr?rr foctidu 17.1 SuperfamiIy Cylindrocorporoidea 180 Family C'\ lindrocorporidae Goodey 180 Genus itivctoluiiinc.s I80 :I c lo/arinuLc 1a1illalrt.s sp. n. 18 :11i ctolainucs oreodoxus sp. n. 187 Biology of Alttlolai,nus oreodoxity sp.n. 193 F.mhryogcncsis 193 Post-embryogenesis 195 Discussion 204 Summar\ 209 1(eI renccs 214 / ice..` l The nematodes are highly diversified lower invertebrates. The name nematode comes from the Greek word 'nematos' which means 'thread' and 'eidos' meaning 'form'. Nematodes are putatively the most species-rich animal phylum. The have various life styles and occur in a variety of habitats, ranging from free-li\ ing in marine, fresh-waters or in soil or parasites of different kinds of plants and animals. I he\ are found in almost all t\ pes of habitats and occur in unimaginable numbers, sires, shapes and structures. Nematoda is a \er\ successful group of animals, four of even five multicellular animals on the planet are nematodes (Gunapala et a/., 1998). They are present almost everywhere such as crop fields, in sand dunes, sediments beneath the ocean floor. freshwaters, in plants, animals and even in humans. This croup is perhaps the most abundant group among multicellular animals on earth. They are di\erse metazoans with an estimated total number of a million species (l.ambshead. 2004). A single acre of soil from the arable lands may contain more than three billion nematodes. while the marine beach sand ma\ contain more than one and a half billion nematodes per acre. Their abundance reaches many million individuals per m- in soil and bottom sediments of aquatic habitats and a handful of soil may contain more than 50 different species of nematodes. Sur e\'s of benthic sediments suggest that the total species number for marine nematodes may exceed one million (1.ambshead. 1993: Lambshead, 2001) with only a f'c~w thousands described in the scientific literature (Malakhov, 1994: De Lc \ & Blaxter. 2002). In terrestrial systems, nematode diversity appears to be under-reported (Lawton et al., 1998), 'ith 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: Boar & Yeatcs, 1998). Their size too is extremely variable ranging from less than I00 fun (GrceJicllu minutu?n) to greater than 8 metres (Pla(enton'!na gigantissimn) discovered in the placenta of a sperm hale. Several Lon i(lHrr►s species exceed 10mm in length. From an environmental point of view. nematodes are part of nearly all ecos'stems in their roles as bacterivores, herbivvores, parasites of animals and plants, and consumers of dissolved as well as particulate organic matter. They have critical roles in flow of energy and cycling of' nutrients. From an anthropocentric point of view, they parasitize humans and plants. domestic and \\ild animals and they can serve as indicators of environmental change. With regard to health, for example. a large proportion (22%) of the human population is parasitized by the intestinal parasite Ascaris• lumbricoules (Crompton, 1988). Considering their impact on crops. McCarter (2009) estimated a global total loss of $118 billion for 2001, of which nearly halt' was related to only two crops. rice and maize. This being so, it is remarkable that they are among the least studied, with only close to 26.000 (< 3% of estimated) species described to date (I lugot etal.. 2001: 1-lallan. 2007). The deepest oceans, freezing polar seas, hot water springs, arid deserts, alpine lakes, mountain tops and the frozen Antarctica, represent the hostile climates of «orid. but surprisingly nematodes are not only present but quite in abundance in all these ecosystems. Tropical rain-forests and also the subtropical areas of the world have tremendous nematode diversity. Soil nematode diversity has been one of the main foci in ecological researches. Nematodes occupy a central position in the soil food %%eb and play significant roles in ecological processes such as nitrogen cycling and plant growth patterns (Neher, 2001). Their abundance, trophic classification and various indices such as Shannon-Weaver index, dominance index, enrichment index, structure index, channel index all have been used to varying degrees to assess soil management (Berkelmans et al.. 2003: Bongers. 1990: Ferris et al., 2001, Yeates, 1999). Soil nematodes offer great potential for use as indicators of biodiversity and 2 ecological stability, and for assessing the impact of changing land use on soil conditions.