E:\ 82(1) 19165-K M Singh and M M Kumawat 2020

Indian Journal of Entomology, 82(1): 000-000 (2020) DoI No.:

ARTHROPOD AND CONSERVATION BIOLOGICAL CONTROL IN

K M Singh and M M Kumawat*

Central Agricultural University, Lamphelpat, Imphal West, Imphal795004, Manipur *College of Agriculture, Agriculture University, Jodhpur- 342 304, Rajasthan *Email: [email protected] (corresponding author)

ABSTRACT

Rice ecosystems have a rich and diverse fauna of which varied in field in different ecological conditions. Beside the phytophages and detritivores, a large number of predators and also exist in the rice ecosystem. Biological control has shown a great potential for reducing the dependence of crop protection on chemical control. Among the predators, spiders are highly abundant in rice fields which feed on variety of pests. The different organisms present in the rice ecosystem are linked by food web which can help insect management because they clarify the functional interaction between populations. Conservation biological control can be achieved by reducing the pesticide-induces mortality of natural enemies and habitat manipulation to improve natural enemies fitness and effectiveness in the rice fields. There is a plethora of natural enemies in rice ecosystem and conservation biological control plays a major role in reducing pests. The identification of safer chemical pesticides and supplemental food of natural enemies is still needed.

Key words: Rice, conservation, biological control, food webs, trophic relationships, natural enemies, insect pests, habitat manipulation Rice ( sativa L.) is one of the most important genetic treadmill beside the treadmill. In crops in the world. It is the primary staple food for recent decades, integrated pest management (IPM) has billions of people in Asia and for hundreds of millions shown great potential for reducing the dependence of of people in Africa and Latin America. This annual grass crop protection on chemical control methods (Pretty et is widely cultivated from 50° N to 40°S latitude and al., 1998; Atanassov et al., 2002) and biological control in areas from sea level to altitude of more than 2,500 is a major component of the IPM programs. metre (Tsunoda and Takahashi, 1984). The major rice Biological control may involve the inundative ecosystems are wetland (also known as paddy or low release of large number of bio- control agents or the land), dryland (or upland) and deepwater. Wetland rice inoculative release of exotic agents. The inundative may be irrigated, rain fed and recession rice. Recession release is economically liable only for those natural rice is grown when rice seedlings are transplanted into enemies which can be cultured invitro or those with receding water which occurs when a lake dries up. simple mass production technique like Trichogramma Earth embankments (bunds) are prepared in wetland spp. Further, availability of these bio- control agents rice fields to retain water. In deepwater rice, standing at the farmers label are also a problem as there is water deeper than 50 cm remains in the field for one limited number of natural enemies production units and month or longer during the growing season (Catling, difficult to transport for longer distance as they are the 1992). Rice cultivation of high input responsive living entities. The inoculative release of exotic agents, varieties which necessitate application of increasing otherwise known as classical biological control, carries amounts of agrochemicals, leading to enhanced a risk of introducing that will have severe non- environmental degradation and reliance on high capital target effects (Howarth, 2000; Twyford, 1991). Beside and production inputsPreview (Tilman,1998; Matson et al., these around 90% of arthropods releases for arthopod 1997; Tilman et al., 2002). These escalating usage of pest control, fails to bring the target under effective the , fungicides and herbicides eroded the control (Gurr et al., 2000). In the attempts to introduced on farm biodiversity (Stark and Banks, 2002; Relyea, exotic parasitoids against stem borers in , one was established against pink stem borer 2006) and causes pest resurgence. With development of but no beneficial effect was observed (IRRI, 1971). new biotype of pests, breakdown of resistance becomes To control rice black bug, two egg parasitoids was a common phenomenon and need to replace the new introduced in the Island of Palawan, however, none of resistance varieties after every 2-3 years. This leads to the species was established (Ooi and Shepard,1994). 2 Indian Journal of Entomology, 82(1), 2020

Considering the problem associated with inundative lividipennis Reuter being the most abundant species. and inoculative biological control, from the last decade, Spiders were the next dominant group which includes attention has been drawn toward the conservation Pardosa pseudoannulata (Bosenberg & Strand) and biological control. Conservation biological control is three species of Tetragnatha. defined as ‘modification of the environment or existing practices to protect and enhance specific natural enemies Survey of farmer’s fields in recorded 435 of other organisms to reduce the effect of pests”. Many species belonging to 266 genera and121 techniques that can be successful in rice fields for families in 13 orders (Inthavong et al., 1998). Rai et managing the pests are discussed herein. al., (2000) compiled an inventory of the biodiversity of coleopterans associated with various rice growing Biodiversity in rice ecosystem ecosystems listed 368 species, which have various Rice is grown in different ecological conditions thus kind of associations with rice, both herbivores and have varied community structure of arthropods and even non-herbivorous. Biosystematics of these revealed in the same field change in the environmental condition that there are 368 species under 191 genera and 26 favours another community structure of arthropods. families. The families Chrysomelidae, Coccinellidae, Rice ecosystems also have a rich and diverse fauna of Curculionidae and Scarabaeidae comprised major arthropods. The arthropod community associated with share followed by Anobiidae, Anthicidae, Bostrichidae, irrigated rice in Philippines was studied by Heong et al. Carabidae, Cucujidae, Dascillidae, Dermestidae, (1991). They reported that phytophages and predators Dytiscidae, Elateridae, Heteroceridae, Hydrophilidae, were predominant in all sites. The phytophages were Languriidae, Melyridae, Meloidae, Ptinidae, Silvanidae, mainly Homopteran and dominated by the cicadellids, Staphylinidae, Tenebrionidae and Trogossitidae. An Nephotettix virescens (Distant), Nephotettixnigropictus analysis of their interaction with rice indicated that (Stal), and the delphacids, Nilaparvata lugens (Stal) a great majority of them are pests, either in the field and Sogatella furcifera (Horvath). Predetors were or storage. Many of them were predators, indicating mainly Heteropterans which includes the veliid, their potential in biological control of pests and can be Microvelia douglasis atrolineata Scott, the mesoveliid, included in integrated pest management. The arthropod Mesovelia vittigera Horvath and the miried, Cyrtorhinus biodiversity is increased with the reduction of chemical pesticides in rice ecosystem (Table 1). Table 1. Comparision of arthropod biodiversity after reduction in use of insecticides in International Rice Research Institute farm in 1989 and 2005

Guilds Biodiversity parameters 1989 2005 Herbivores Abundance % 46.2% 11.6%

Species richness, S or Esn (rarefaction) 13.6 36.0 Log series index α 3.10 8.97 Reciprocal Simpson’s (I/D) 2.25 2.56

Exp Shannon or Hill N1 3.46 5.75 Predators Abundance % 40.0% 58.0%

Species richness, S or Esn (rarefaction) 37.6 65.0 Log series index α 6.38 12.28 Reciprocal Simpson’s (I/D) 5.12 6.50

Exp Shannon or Hill N1 8.25 11.70 Parasitoids Abundance % 5.6% 4.3%

Species richness, S or Esn (rarefaction) 17.1 38.0 Log series index α 5.41 14.67 PreviewReciprocal Simpson’s (I/D) 2.57 13.25 Exp Shannon or Hill N1 5.37 20.91 Detritivores Abundance % 8.1% 26.1%

Species richness, S or Esn (rarefaction) 5.6 30.0 Log series index α 0.88 5.70 Reciprocal Simpson’s (I/D) 1.19 8.02

Exp Shannon or Hill N1 1.46 10.80 Source: Heong et al., 2007 Arthropod biodiversity and conservation biological control in rice 3 K M Singh and M M Kumawat

Insect pests predators inhabiting in the paddy fields. They prey Rice is primarily a crop of warm and humid on wide range of insect viz. stem borers, leaf folders, environments where the insect pest problems are very plant hoppers, leaf hoppers and some dipterans. So, severe and numerous. A good attempt has been made by they are one of the generalist predators. Barrion and a number of workers to study the insect pest complex Litsinger (1984) has made a good compilation of field associated with rice crop. Grist and Lever (1969) spiders from 12 Asian countries, listing 173 species listed over 800 species of damaging rice, either from the temperate zone (Japan, , and in standing crop or in storage. Pawer (1974) listed ) as compare to 86 species from the tropics 650 species of insect pest collected from rice field in (, Burma, , , , Philippines. Pathak (1977) mention about 100 species of Philippines, Srilanka and ). Xu et al., (1987) insect attacking rice of which about 20 are major pests. also reported 167 species falling to 28 families from the Further, about 159 pest species of rice were identified rice ecosystem of Zhejinag, China. Surveys in different by Rob (1990) from Bangladesh. part of the northern and western India have revealed that there are more than 20 species of spider which suppress The diversity of pests in the rice ecosystem the population of brown plant hopper, white backed depends on the environment. The distribution of the plant hopper, green leaf hopper, leaf folder and whorl monophagous species, Asian rice gall midge, Orseolia maggot in rice ecosystem (Bhathal and Dhaliwal, 1990). oryzae (Wood-Mason) coincides with the distribution Thirty-four species of spider belonging to 10 different of wild rice varieties. Armyworms, Mythimna separate families from Cuttack, Orissa and 5 species from Assam Walker and cutworms, Spodoptera litura (Fab.) attack the have also been reported by Rai and Chandrasekar (1979) pre-flooded crop. As the fields flooded, aquatic species and Krishnaswamy et al., (1984), respectively. In central enter the ecosystem such as black bugs, Scotinaphora India, 13 species of spiders belonging to 8 families were spp. and rice caseworm Nymphula depunctalis (Guenee). found to be common from the nursery stage to harvest Thrips, Stenchaetothrips biformis (Bagnall) and (Bhardwaj and Pawar, 1987). mealybugs, Brevennia rehi(Lindinger) thrive in rice during droughts. In deepwater rice, yellow stem borer Beside spiders, several hemipteran predators feeding (YSB), incertulas Walker and rice hispa, on various insect pests of rice were also reported from Dicladispa armigera (Olivier) are dominant. In dryland different regions. Cyrtorhinus lividipennis Reuter and rice fields, ants, whitegrubs, black , termites, Tytthus sp. are important predators of plant hoppers field crickets, mole crickets, root aphids, mealy bugs, and in different rice growing areas root bugs, seedling maggots, grasshoppers, thrips and (Balasubramanian et al., 1988; Basilio and Heong, seed bugs cause damages. Rice crop grown near maize 1990; Peter, 1988). A large population of Hydrometra is prone to Chilo stem borer infestations and rice grown albolineata (Scott) feeding on plant and leaf hoppers of near sugarcane is prone to pink stem borer, Sesamia rice was observed in Hunan, China (Wu, 1988). Water inferens (Walker) infestations. Grasslands near rice strider Limnogonus fossarum (F.) is an effective predator fields can serve as a source of armyworms, grasshoppers, of leaf hopper, plant hopper and other pests is present whitegrubs and seedling maggots (Jahn et al., 2007). in different water bodies including wetland rice(Rubia Changes in the environment may favour one pest groups and Heong, 1990). The hemipteranpentatomid but not another (Loevinsohn, 1997). Flooding a rice field Amyotea (=Asopus) malabarica (Fab.) predates on eliminates soil pests whereas aquatic species thrive in the larvae of Melanitis ledaismene (Cramer), Parnara ponded conditions. Prolonged droughts in mid-1980s (=Pelopidas) mathias (Fab.), P. naso (Fab.), Spodoptera in Indonesia and the southern Philipppines (Glanz, spp. ,Mythimna loreyi Duponchel, Mythimna venalba 1996) led to a shift in stem borer species from the non (Moore), Psalispennatula (Fab.), Laelia fasciata aestivating YSB to the aestivating white stem borer (Moore), Euproctis xanthorhoea (Koller), Chilo spp., (WSB), Scirpophaga innotataPreview (Walker). S. incertulas and S. inferens (Pati and Mathur, 1986). A pyrrhocorid bug, Antilochus nigripes (Burm.) was found Predators feeding on the nymphs and adults of Leptocorisaacuta Among the predators, spiders are highly abundant Thumb in Eastern India. Its predation ranged from 1.66 in rice fields preying on a wide array of insect pests to 5. 66 bugs per day (Rao and Prakash, 1999). and have a major role to play in the natural regulation Many orthopteran and coleopteran predators were of insect pests. Spiders have higher host finding ability also recorded in the rice ecosystem. The katydids, and greater feeding potential in comparison to other 4 Indian Journal of Entomology, 82(1), 2020

Conocephalus longipennis (Haan) and Conocephalus and pupal parasitoids of the lepidopteran pests of rice maculates (Le Guillou) predates on the egg masses of were also reported. lepidopteran pests of rice, leafhoppers, plant hoppers, early instars nymphs of grasshoppers and eggs of A total of 61 species of hymenopterans was identified oratorius (Fab) (Deng and Jin, 1985; De as parasitoid of C. suppresalis in Spain, 22 of which was Kraker et al., 2001). However, after flowering, the ichneumonids, 25 braconids and 14 chalcids (Jimenez katydids feed on flowering and later on young grains et al., 1992). The braconid, Apantales cypris Nixon (Manley, 1985). The , Metioche vittaticollis parasitized the of C. medinalis in China with an (Stal) prayed on the eggs of a range of insect average parasitism of 28.6% (Wei, 1987). In rice fields, pests of rice including Chilo suppressalis (Walker), the braconid Apentalis ruficrus (Haliday) gave good Cnaphalocrosis medinales (Guenee), Hydrellia control of Mythimna separata in late March- April and philippina Ferino, M. separata and to a lesser extent of Naranga aenescens Mooreand the hesperiid Parnara L. oratorius (Rubia and Shepard, 1987; Lee et al., guttatus (Bremer & Grey) in late July to Mid August in 1997). Shepard and Rapusas (1989), Chander and China (Zhang, 1986). A torymid parasitoid, Podagrion Singh(2003) observed Paederus fuscipes Curtis, ahlonei Mani & Kaul was recorded for the first time Ophionia indica(Thunberg), Verania (=Micraspis) in India as a parasitoid of C. medinalis (Kaul, 1993). discolor (Fab), Micraspis crocea (Mulsant), Coccinella Aulosaphes sp. was recorded as the dominant parasitoid arcuata Fab (=Harmoniaoctomaculata), Coccinella of C. medinalis followed by Bracon sp. in Assam. Other septempunctata Linn, Brumus (=Brumoides) suturalis parasitoids recorded were Gonozius sp., Cardiochiles (Fab), Menochilus (=Cheilomenes) sexmaculatus philippennis Ashmead and Temelucha (=Temeluchella) (Fab) and Coccinella transversalis Fabas conspicuous sp. (Borah and Saharia, 1989). Mishra and Senapati predators in rice fields. Kobayashi et al., 1990 reported (1996) reported Goniozus triangulifer Kieffer as a larval carabid, Ophionia indica which predate on Orseolia parasitoid of C. madinalis. Brachymeria lasus (Walker) oryzae. Many dragon and damsel flies were also was reported as a pupal parasitoid of Mythimna separata observed on the rice ecosystem by Garg et al. (2002). (Husni et al., 2001). In Uttar Pradesh, India, Apentalis (Cotesia) ruficus (Haliday) was found to cause larval Parasitoids mortality of M. separata upto 75.35% during 1996 Dey et al. (1999) published a checklist of (Pandey and Sharma, 1999). hymenopterous parasitoids associated with rice Against leafhoppers and plant hoppers, Huang agroecosystem. In China and Philippines, Trichogramma (1984) reported Pseudogonatopus flavifemur (Esaki & spp. plays a vital role in reducing the population of Hashimeto) as an important biocontrol agent of brown Marasmia patnalis Bradley, C. suppressalis and C. plant hopper (BPH) in China. The young stage of the medinalis (Zhou, 1986, Arida and Shaperd, 1990). In hymenopteran parasitized the pest and adult females Java, the scelionid, Telenomus rowani (Gahan) was prayed on nymphs of the pest. Anagrus flaveolus the major parasitoid species followed by the eulophid, Waterhouse (: Mymaridae) is an important Tetrastichusschoenobii Ferriere and Trichogramma egg parasitoid of BPH in Philippines (Chandra and japonicum Ashmead (Soejitno, 1988). She and He Dyck, 1988) and P. flavifemur a nymphal parasitoid of (1988) revealed the presence of 19 parasitoid species BPH in Korea Republic (Kim et al., 1988). Gonatocerus attacking the pyralid C. suppressalis. The dominant cincticipitis Sahad and Paracentrobiaandoi (Ishii) was species were T. japonicum, Chelonus munakatae reported as egg parasitoid of the green leaf hopper, N. Matsumura, Microgaster russata Hal., Centeterus cincticeps in Japan (Miura, 1990a and Miura, 1990b). In alternecoloratus Cush. (Auberteterus alternecoloratus) , N. lugens were parasitized by the mymarids and Eriborus terebrans (=Diadegma terebrans) Anagrus sp. nr flaveolus and A. optabilis and the (Gravenhorst). MahrubPreview (1993) identified three species trichogrammatid, Oligosita sp. Nephotettix spp. eggs of egg parasitoid of S. incertulas from Indonesia. were parasitized by the two species of the mymarid They were T. schoenobii, T. rowani and T. japonicum. Gonatocerus and one trichogrammatid genus In Orissa (India), S. incertulus eggs were parasitized Paracentrobia. Gonotocerus spp. and Paracentrobia by Telenomus dignoides (Nixon), T. japonicum and sp. seldom attacked the same Nephotettix egg T. schoenobii (Ram et al., 1996) and in Punjab, batch (Fowler et al., 1991). In India, Ecthrodelphax the scelionid, T. dignoides parasitized the egg of S. fairchildii (Perkins), Haplogonatopus apicalis Perkins, incertulas. Beside the egg parasitoids several larval Pseudogonatopus hospes (Perkins) and a species of the Arthropod biodiversity and conservation biological control in rice 5 K M Singh and M M Kumawat genus Pseudogonatopus near P. pusanus were recorded O. oryzae (Mathur et al., 1988). Rana et al., (2002) attacking N. lugens and S. furcifera (Yadav and Pawar, reported that parasitism of P. oryzae reaches upto 42% 1989). with 20.05 to 25.05% silver shoot. In Manipur (India), P. oryzae is the predominant parasitoid of O. oryzae and Morrill and Almazan (1990) reported Gryon the parasitism even reach upto 69.5% (Singh, 2003). nixoni Masner (=G. flavipes) as an egg parasitoid of L. oratorius. The Malayan black bug, Scotinophara Food webs coarctata (Fab) in Palawan Island, Philippines was Most of the works on the biological control of attacked by the scelionid egg parasitoid, Telenomus insect pests of rice are confined on the survey of triptus (Nixon) which parasitized a mean of 61.4% of natural enemies, their extent of predation or parasitism the egg masses and a mean of 19.2% of the eggs (Perezet and promoting one or two potential natural enemies. al., 1989). In Bangladesh, rice hispa, Dicladispa However, in the rice field hundreds to thousands of armigera (Olivier) grubs were parasitized by braconid different species of plants, insects, arachnids, fungi, and eulophid parasitoids (Islam and Rabbi, 1998). nematodes, snails, vertebrates and other organisms are Trichogramma zahiri Polaszek was reported as an egg present and they are linked by a vast, intrinsic network parasitoid of the rice hispa, D. armigera in Bangladesh of feeding relations called food web. According to (Polaszek et al., 2002). Cohen et al., (1994), food webs can help insect pest There are also some reports on the parasitoids of management because they clarify the functional the cecidomyiid pest, Orseolia oryzae. In Sri Lanka, interaction between populations (Fig. 1). Settle et al., five hymenopterous parasitoids were found parasitizing (1996) had the opinion that consistently high level O. oryza. They were Platygaster oryzae Cameron, of natural biological control may often result from a Platygaster foersteri (Gahan), Neanastatus cinctiventris complex set of community-level interactions that lead Girault, Obtusiclava oryzae Subba Rao and Eurytoma to a far more stable and robust system. But only in some sp. P. oryzae parasitized upto 13.5% followed by N. cases, agricultural data on pests and their biological cinctiventris (8.7%) (Kobayashi et al., 1990). In Orissa control, species have been organized into food webs. (India), the egg larval parasitoid P. oryzae was found Since 1977, entomologists of the International Rice to be more active against the rice pest O. oryzae and Research Institute (IRRI) have been describing the food the larval and pupal parasitoid Propicroscytus mirificus web of Philippines irrigated and rainfed rice fields. The (Girault), Eurytoma spp. and Neanastatus grallarius whorl maggot food web comprises 114 taxa including Masi (=N. cinctiventris) were more active against the 71 predators, 13 parasitoids and 62 secondary natural grass midge, Orseolia mnesitheae Gagne (Mathur et enemies (IRRI, 1987). The food web of semi-aquatic al., 1991). A solitary Platygaster sp. also parasitized Preview

Fig. 1. Trophic relationships in a rice ecosystem (Source: Heong, 2012) 6 Indian Journal of Entomology, 82(1), 2020 rice caseworm consists of 99 taxa with 78 predators, outbreak of N. lugens is associated with insecticide only 4 parasitoids, but 39 secondary natural enemies usage not with fertilizers (Den Braber and Meenankanit, (IRRI, 1990). However, no egg parasitoid was recorded 1992). Field experiments had shown that insecticide against the pest. It may be due to its egg laying site. sprays destroyed natural enemies (Heinrichs, 1994), Among the stem borers, yellow stem borer, Scirpophaga destroyed detritivores (Settle et al., 1996), disorganized incertulas comprises 118 species including 57 predators, predator-prey relationship and food chain linkages 38 parasitoids and 23 secondary natural enemies (IRRI, (Cohen et al., 1994; Schoenly et al., 1996) and 1984) and Chilo stem borers includes 111 species with favoured the development of secondary pests, such as 57 as predators, 33 parasitoids, 18 hyperparasitoids plant hoppers (Heinrichs and Mochida, 1984; Heong and 28 secondary predators (IRRI, 1989). Leaf folders and Schoenly, 1998). Further, many researchers even are a group that is heavily attacked by natural enemies hypothesized that heavy use of insecticides actually (Khan et al., 1988). The food web from the Philippines accelerates the adaptation of the pest populations to the comprises 163 taxa with 77 predators, 51 parasitoids resistant varieties (Settle et al., 1996). and 24 secondary natural enemies. Leafhoppers are under heavy pressure from natural enemies beginning Competition with weeds and adverse abiotic factors, with trichogrammatids (Paracentrobia, Oligosita) and especially inadequate water, can decrease host-plant mymarids (Gonatocerus, Anagrus) egg parasitods resistance (Litsinger, 1993) and also diminish ability (Jahn et al., 2007). Predators alone reduce the leaf to compensate the injury caused by the pests. Survey folder number by 50-70% in field studies (Ooi and in irrigated fields of Philippines indicated that high Shepard, 1994).The food web of the Nephotettix spp. yields are strongly correlated with good overall crop includes 205 taxa with 129 predators, 35 parasitoids, husbandry but not with the use of insecticides (Way and 6 pathogens and 63 secondary natural enemies (IRRI, Heong, 1994). Dead heart cause by the stem borers in 1986). Of the brown plant hopper food web, only 11 the early crop stage does not affect much on the yield of the 76 taxa are parasitoids and 11 secondary natural as the rice plant can recover the injury. Potentially enemies and the remaining are predators dominated by the most damaging stem borer Scirpophaga innotata spiders (IRRI, 1980). Schoenly et al., (1996) mentioned (Walker)causing whiteheads begins oviposition at the cumulative version of food web of Philippines rice about 80 days after transplanting. However, during ecosystem contains taxa which includes pathogens, this time, a rich natural enemy community has already nematodes, mites, spiders, insects, snails, vertebrates accumulated including the parasitoids and can make and over 10,000 trophic (consumer-resource) links. a major contribution in stem borer control, unless insecticides are used(Way and Heong, 1994). As an Conservation biological control example, when carbofuran was applied for controlling The arthropod diversity and abundance are Scirpophagainnotata, egg predation and parasitism fundamental components of rice ecosystems that have of the pest was reduced to halve and the number of resistance and resilience to pest attacks. They provide whiteheads was doubled (Triwidodo et al., 1992). farms with ecosystem services, such as resistance to In the organic matter rich fields of rice, there is pest invasions and regulation of pest populations that abundant population of detritus feeding and plankton prevent pest species from increasing to levels that feeding insects and they serve as alternative prey for can cause economic loss to farmers. The impetus for many generalist predators (Fig. 2). These alternative better understanding of the role of natural enemies preys can give the predator population a ‘head start’ stemmed from widespread and devastating outbreaks on later-developing pest populations. This process of Nilaparvata lugens associated with early Green can strongly suppress pest populations and generally Revolution Technology in Tropical Asia (Way and lend stability to rice ecosystems by decoupling Heong, 1994). At first,Preview the tillering and other growth predator populations from a dependence on herbivore characteristics were largely blamed for the outbreaks populations (Settle et al., 1996). Feeding studies on (Dyck et al., 1979). However, such outbreaks were Atypena formosana (Oi) by Sigsgaard et al. (2001) also associated with the widespread and intensive used suggest that alternative prey such as collembolans may of certain insecticides which are highly destructive still be more profoundly important. Spider feeding only to natural enemies. In Thailand, despite intensive N. lugens or N. virescens led to very poor survival of insecticide usage, outbreaks of N. lugens became spiders to the adult stage. In contrast, a mixed diet of increasingly serious upto 1989-90 and it confirm the either hemipteran with collembolans and drosophila as Arthropod biodiversity and conservation biological control in rice 7 K M Singh and M M Kumawat

Fig. 2. Number of detritivores and plankton feeders in Indonesia rice fields (Source: Settle et al., 1996) gave faster development time and greater survival. This dry-season crops such as soybeans or green manures illustrated that availability of prey such as collembolan or promoting the conservation of straw mulch piles. is essential for the performance of this linyphiid, not simply an early-season alternative food resource. Beside The majority of the rice pests are specific or the rice pests, some of the insect in the rice ecosystem narrowly oligophagous for rice, so there is little chance feeding on wild plants also serves as an alternative of harbouring rice pests by the plants surrounding host of the many parasitoids and predators. The eggs of the rice fields except some polyphagous pests like Notiphila latigenis Hendel and Notiphila similis Meijere armyworms and locust. On the other hand, non-rice which were usually laid on the stem of perennial fern habitats, particularly the narrow vegetation-covered Marsilea minuta L. and it serve as alternate host of bunds surrounding each field, seem especially the stem borer parasitoid Trichogramma (Barrion and important as a source of natural enemies, particularly Litsinger, 1986). early arriving species such as spiders. The rice bunds are also abundantly colonized by fireants, Selenopsis Settle et al. (1996) mentioned two ways for geminate (F.) and this ant is potentially important disrupting high levels of biological control in the rice predator of N. lugens and other rice pests including ecosystem of Java, Indonesia: corn rootworm, Diabrotica adelpha Harold (Way et al., 2002). Parasitoids of the rice hoppers commonly Early season application of pesticides including parasitized non-rice hoppers species on wild host plants herbicides keeps down the natural enemies’ population. during off season (Way and Heong, 1994). Another Insecticide applications are the most important factor important very early arriving predator of plant hoppers, causing pest problem. Even the herbicides give negative Cyrtorhinus lividipennis Reuter can also survive in effects to phytoplanktons and zooplanktons, which give the off-season as an egg predator of insects on wild indirect negative effect to the predator population. plants (Bentur and Kalode, 1987). Beside these non- Unfavourable landscape design and water use rice habitats seem likely to be an important source of patterns also weaken the system. In rice landscapes some other early arriving predators like that characterized by synchronous and large scale planting can seasonally concentrate on rice (Way and Heong, patterns and precededPreview by long, dry fallow periods, 1994). Drechsler and Settle (2001) reported that high the arrival of natural enemies is severely delayed. A proportion of vegetable fields in the landscape reduces secondary but important effect of this delay is that pest abundance in the rice field. According to Gurr generalist predators could not take the advantage of (2009) fruit and timber trees, may serve as sources and the peak population of alternative prey provided by the sinks for natural enemies. Proximity of such landscape detritivores and plankton feeders due to late arrival. The elements is important because many of the generalist long dry fallow periods can be shortened by planting predators like wolf and web-building spiders, colonized rice from adjacent habitats. 8 Indian Journal of Entomology, 82(1), 2020

Different habitat manipulation techniques for through detritivore prey- conservation of arthropod natural enemies were also developed in different parts of the world. Shepard et al. Enrichment of organic matter of the soil will increase (1989) suggested for placing rice straw bundles in the the population of detritivores and plankton feeders and rice fields during crop free period. The bundles can be it serve as the alternate prey of the generalist predators. prepared by arranging rice straw into cone-shaped tents At the early crop stage, detritivores and plankton about 30cm in diameter at the bottom which serves as feeders are vitally important and need to be conserved. habitat for the predators and prey communities during Application of pesticides reduce their population. crop free period. For management of insect pests of rice, Plank 3: Enhancement of specialist natural enemies straw bundles@ 20 bundles/ha can be placed in the rice by habitat manipulation- fields. The bundles can be first put in sorghum fields for about 15 days to charge the bundles with natural The third plank targets at providing resources, enemies like spiders and rove beetles and then transfer shelter and food for the natural enemies, especially the it to the rice field after 15 days of rice transplanting. specialist. This may be achieved by populating bund While placing the straw bundles, it can be fixed with and non-rice habitats with nectar rich flowering plants. bamboo sticks and the lower portion of the bundle In China when nectar rich sesame plants are grown on should remained 15 cm above the water level. This bunds, parasitoid abundance increased. Sesame flowers technique could increase the spider population in rice also enhance the host searching efficiency by the ecosystem (Tanwar et al., 2011). parasitoids. In , flowers on the bunds increased egg parasitism of plant hopper eggs. In Thailand, In Japan, mowing of the ridges in the rice fields and parasitoid species richness increased when fields are cultivation of Chinese milk vetch, Astragalus sinicus surrounded by flowering plants. In Hainan, Philippines, L. as a green manure crop before rice transplanting fields with clean bunds had lower spider biodiversity. increased the number of wolf spiders in rice field (www. ipmcentres.org, 2012). Placement of weed residues in An alternative approach for manipulating natural upland rice in piles harvour more spider population enemy movement based on chemical ecology may (Afun et al. 1999). Carabid beetles were significantly be viable in the future. Natural enemies are attracted more abundant in plots with strips or piles of weeds to the volatile chemical produce by the plants that and staphylinid beetles were most abundant in mulched are attacked by arthropod herbivores. Some such plots. The rice ratoons were found to be a favourable hervivore-induced plant volatiles (HIPVs) have been habitat for natural enemies, especially predators identified, synthesized and used in slow-released and flooding a field continuously favored predator dispensers or sprays. Under field conditions, methyl- population (Dela Cruz and Litsinger, 1988). Allowing salicylate, cis-3-hexen-1-ol, (Z)-3-hexenyl acetate, the growth of weeds in the roundabouts of small dikes and benzal dehyde have resulted in elevated catches of in rice fields, especiallyPaspalum sp., Digitaria sp. and biological control agents (James, 2005). Application Echinochloa sp. Application of host extract and sugar of jasmonic acid on rice increased the parasitism of N. liquid on rice plant was recommended for management lugens eggs by Anagrus nilaparvatae Pang & Wang of brown plant hopper (Atmaja and Laba, 2000). (Gurr, 2009). Exogenus applications of jasmonic acid to rice plants elevated the levels of several volatiles, Heong (2012) suggested three planks for ecological including aliphatic aldehydes, alcohols, monoterpenes, engineering to enhance natural biodiversity in the rice sequiterpenes, methyl salicylate and n-heptadecane. ecosystem- Plants treated withsuch chemicals not only producing Plank 1: Reduction in insecticide use especially in HIPVs which attract the parasitoids and predators, it the early crop stage- may also affect the pest behavior, making treated plants Previewless attractive to the pests (Karban and Chen, 2007). Application of insecticides in the first 40 days after sowing do more harm than by destroying Conclusions natural enemies and aquatic detritivore prey fauna. Biological control as a main component of integrated It significantly decrease food chain lengths and pest management (IPM) has shown a great potential for disorganize predator-prey relationship. reducing the dependence of drop protection on chemical control methods. Biocontrol approaches like classical Plank 2: Enhancement of generalist natural enemies biological control and inundative and inoculative Arthropod biodiversity and conservation biological control in rice 9 K M Singh and M M Kumawat

release of natural enemies have certain limitations. yield in upland rice in Cate d’lvoire. Biological Agriculture and Considering the problem associated with other Horticulture 17(1): 47-58. biocontrol approaches, from the last decade, attention Arida G S, Shepard B M. 1990. Parasitism and predation of rice leaffolders, Marasmia patnalis (Broadley) and Cnaphalocrosis has been drawn toward the conservation biological medinalis (Guenee) (: Pyralidae) in Liguna Province, control. Rice ecosystems have a rich and diverse Philippines. Journal of Agricultural Entomology 7(2): 113-118. fauna of arthropods. As the crop is grown in different Atanassov A, Shearer P W, Hemilton G, Polk D. 2002. Development and ecological conditions thus have varied community implementation of a reduced risk peach arthropod management structure of arthropods. Beside the phytophages and programme in New Jersey. 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(Manuscript Received: October, 2019; Revised: December, 2019; Accepted: January, 2020; Online Published: January, 2020)

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