The Sugarcane Woolly Aphid, Ceratovacuna Lanigera Zehntner (Hemiptera: Aphididae): Its Biology, Pest Status and Control

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65. Scharff, C., Chasing fate and function of new neurons in adult 75. Nordeen, E. J. and Nordeen, K. W., Sex and regional differences brains. Curr. Opin. Neurobiol., 2000, 10, 774–783. in the incorporation of neurons born during song learning in zebra 66. Nottebohm, F., Why are some neurons replaced in adult brain? J. finches. J. Neurosci., 1988, 8, 2869–2874. Neurosci., 2002, 22, 624–628. 76. Nordeen, K. W. and Nordeen, E. J., Projection neurons within a vocal 67. Gross, C. G., Neurogenesis in the adult brain: death of a dogma. motor pathway are born during song learning in zebra finches. Nature, Nature Rev. Neurosci., 2000, 1, 67–73. 1988, 334, 149–151. 68. Kempermann, G., Why new neurons? Possible functions for adult 77. Nadel, L., The hippocampus and space revisited. Hippocampus, hippocampal neurogenesis. J. Neurosci., 2002, 22, 635–638. 1991, 1, 221–229. 69. Barinaga, M., Newborn neurons search for meaning. Science, 2003, 78. Bischof, H.-J., Environmental influences on early development: a 299, 32–34. comparison of imprinting and cortical plasticity. In Perspectives in 70. Bolhuis, J. J. and Macphail, E. M., A critique of the neuroecology Ethology (eds Bateson, P. and Klopfer, P.), Plenum Press, New York, of learning and memory. TICS, 2001, 5, 426–433. 1985, vol. 6, pp. 169–217. 71. Shingo, T. et al., Pregnancy-stimulated neurogenesis in the adult 79. LeVay, S., Wiesel, T. N. and Hubel, D. H., The development of ocu- female forebrain mediated by prolactin. Science, 2003, 299, 117– lar dominance columns in normal and visually deprived monkeys. J. 120. Comp. Neurol., 1980, 191, 1–51. 72. Eichenbaum, H., To cortex: thanks for the memories. Neuron, 1997, 80. Wiesel, T. N. and Hubel, D. H., Ordered arrangement of orientation 19, 481–484. columns in monkeys lacking visual experience. J. Comp. Neurol., 73. Tramontin, A. D. and Brenowitz, E. A., Seasonal plasticity in the 1974, 158, 307–318. adult brain. Trends Neurosci., 2000, 23, 251–258. 74. Lee, D. W., Smith, G. T., Tramontin, A. D., Soma, K. K., Brenowitz, ACKNOWLEDGEMENT. Grant support from DST-SERC for young E. A. and Clayton, N. S., Hippocampal volume does not changes scientists (NO. SR/FTP/LS-192/2000), New Delhi is acknowledged. seasonally in a non food-storing songbird. Neuroreport, 2001, 12, 1925–1928. Received 23 May 2003; revised accepted 9 March 2004

The sugarcane woolly aphid, Ceratovacuna lanigera Zehntner (Hemiptera: Aphididae): its biology, pest status and control

Sunil Joshi* and C. A. Viraktamath Department of Entomology, University of Agricultural Sciences, GKVK, Bangalore 560 065, India

Sugarcane woolly aphid, Ceratovacuna lanigera Zehnt- Recent surveys in the pest-affected areas of Mahara- ner has been recently reported in outbreak propor- shtra (western India) and Karnataka (southern India) tions from western and southern India. Though the have yielded several indigenous predators which in- pest was first reported from West Bengal in 1958 and clude coccinellids, neuropterans, syrphids and a pyra- later from other parts of Northeast India, it had not lid. Integrated pest management involving mainly made its way to other parts of India. The pest breeds mechanical and biological control appears to be the on plants of the family Poaceae, but has been also best option. There is ample scope for more directed observed on members of Bixaceae, Theaceae and Com- work on this important emerging pest, especially in bretaceae. It has been recorded on ten species of plants the areas of pest ecology and distribution. The role of in India. It reproduces parthenogenetically and has an resistant varieties, and biological, cultural and mecha- anholocyclic (absence of sexually producing genera- nical control in managing the pest needs to be care- tion) life cycle. Thirty-eight natural enemies have been fully evaluated. recorded on the aphid from different parts of the world.

SUGARCANE is attacked by several aphid species in India. three to Hormaphidinae. Among the three hormaphidine Raychaudhuri1 listed 17 species of aphids associated with species in the genus Ceratovacuna Zehntner, Ceratova- sugarcane of which seven belong to subfamily Aphidi- cuna lanigera Zehntner is a serious pest of sugarcane in nae; five to Pemphiginae; two to Drepanosiphinae and several parts of the Oriental region. The species is known from India, Nepal, Bangladesh, East and South Asia, Fiji and Solomon Islands2 (Table 1). It has been recorded on *For correspondence. (e-mail: [email protected]) other members of the family Poaceae like bamboo and

CURRENT SCIENCE, VOL. 87, NO. 3, 10 AUGUST 2004 307 REVIEW ARTICLES other grasses. C. lanigera lives in large colonies, sucking breed on plants belonging to the families Styracaceae, phloem sap from the leaves and excreting copious honey- Poaceae, Arecaceae and Orchidaceae. dew onto foliage leading to the development of sooty Among the 14 species of Ceratovacuna known from the mould. The direct and indirect damage affects sugarcane world4, seven occur in the Indian region, viz. silvestrii yield and quality. In India, the pest was known only from Takahashi, perglandulosa Basu, Ghosh and Raychaudhuri, Northeast India, but recently, it has made its entry into indica Ghosh, Pal and Raychaudhuri, spinulosa Ghosh western and southern parts of the country. This article and Raychaudhuri, lanigera Zehntner, graminum van der reviews the present status of knowledge on the biology, Goot and nekoashi (Sasaki). Among these, indica, per- behaviour, pest status and control strategies of the pest. glandulosa and spinulosa are endemic and are known only by apterae viviparae. C. nekoashi (Sasaki) is known only from northwest Himalaya4, species, viz. silvestrii, Taxonomy and nomenclature perglandulosa, indica, spinulosa, lanigera and nekoashi are known from northeast India. Patil5 reported the pre- In 1897 Zehntner erected the genus Ceratovacuna under sence of C. graminum van der Goot in India. The US the tribe Cerataphidini with C. lanigera Zehntner as the National Collection of Aphididae records also indicate type species. The name C. lanigera is preferred to Oregma deposition of a specimen of C. graminum van der Goot of lanigera van Deventer (1906) and Cerataphis sacchari- Indian origin. However, there is no literature indicating the vora Matsumura (1917), which are junior synonyms of occurrence and distribution of this species from south or this species3. This genus is characterized by the presence north India. Among all the species known from India, the of frontal processes in apterae and alatae. Apterae often most commonly distributed species is C. lanigera. possess crenulated margins of wax glands arranged in a row up to the seventh and eighth tergites, with similar wax glands on the lateral margin. Such wax glands are Present status of the pest in India absent in alatae. Head in apterae is fused with prothorax or with entire thorax, and the eighth tergite is always The pest was first recorded in West Bengal6 and later in free. Forewings are with media once branched and hind- different parts of Northeast India as a minor pest. In Maha- wings with two oblique veins. The species of the genus rashtra, incidence of C. lanigera was recorded for the first time in July 2002 in Sangli district. Later, the infestation spread to Kolhapur, Satara, Pune, Solapur and Table 1. Geographical distribution of Ceratovacuna lanigera82 and Ahmednagar. A total of 15.5% of the area under sugar- sequence of its appearance in different states of India cane is presently infested, with the highest infestation in

Place Pest status Year of appearance Reference Sangli (25.29% of total sugarcane area) followed by Kolha- 7 pur (21.27% of total area) . Brunei Darussalam Minor 1993 84 In Karnataka, the infestation was first observed in Fujian Minor 1927 85 Guangdong Minor 1928 86 Athani taluk, Belgaum district in September 2002. Sub- Guangxi Minor 1945 87 sequently, it was recorded in Raibag, Chikkodi, Hukkeri Taiwan Major 1910 88 and Gokak taluks. So far, infestation has been recorded in Yunnan Minor 1928 86 Dutch East Indies Minor 1925 89 five districts, viz. Bagalkot, Bijapur, Bellary, Bidar and Indonesia Minor 1993 84 Belgaum. The highest area of incidence was recorded in Java Major 1900 90 Belgaum with 51.55% of total sugarcane area infested, Kyushu Minor 1924 91 Ryukyu Archipelago Major 1971 92 followed by Bagalkot with 31.90%. Incidence in Bidar Malaysia Major 1977 93 was the lowest (0.43% of total sugarcane area)8. The in- Myanmar Minor 1935 94 festation has now spread to other districts of Karnataka, Philippines Major 1917 95 Sri Lanka Minor 1971 92 viz. Koppala, Davangere, Haveri, Shimoga and Raichur Thailand Minor 1993 84 (through results of survey conducted by C.A.V.). Vietnam Major 1928 86 Apart from Maharashtra and Karnataka, the woolly aphid Fiji Minor 1946 96 Papua New Guinea Minor 1953 97 has spread to Uttar Pradesh, Andhra Pradesh (pers. commun. Indo-China Minor 1941 98 RRS, Rudruru and RRS, Vuyyuru), Bihar and Uttaranchal. Recently, attempts were made to predict the potential India West Bengal Minor 1958 6 geographic spread of the pest by using two computational Tripura Minor 1963 99 approaches, viz. GARP and DIVA-GIS9. The studies indi- Assam Minor 1967 100 cated that the pest has low probability of spreading to Uttar Pradesh Minor 1971 101 Arunachal Pradesh Minor 1973 102 eastern parts of Maharashtra and Andhra Pradesh and is Sikkim Minor 1979 103 not likely to spread to Tamil Nadu, Kerala and the coastal Maharashtra Major 2002 5 areas. However, in Karnataka the pest is likely to occur Karnataka Major 2002 104

along the transitional belt and in the northern and sou- 308 CURRENT SCIENCE, VOL. 87, NO. 3, 10 AUGUST 2004 REVIEW ARTICLES thern dry zones, but is less likely to occur in south inte- days3 and 32–40 days14. The average longevity of apter- rior Karnataka. ous and alatae adults was 36 and 8.3 days12, 20.5 and Recent surveys have indicated invasion of the pest in 24.1 days13. Average fecundity of apterous adult was 60 the villages of Vadakali, Lingapura and Anegula, Kekkeri aphids, while it was 10 aphids in alatae12. The rate of re- taluk, Mandya district and Doddamalugu, T. Narasipura production under laboratory conditions varied from 3 to 5 taluk, Mysore district. Apart from Mandya and Mysore of nymphs per day, with total fecundity of 41 to 56.6 aphids13. Karnataka, the pest has recently entered into Tamil Nadu In Philippines, sugarcane varieties were found to influ- and Kerala also. ence the life span and number of progeny produced by the Keeping in view the seriousness of sugarcane woolly aphid15. Longer nymphal duration and lower fecundity aphid infestation, the Department of Agriculture and Co- were recorded on resistant varieties16. operation, Ministry of Agriculture, Government of India has constituted a central team, which is presently investi- gating the causes of outbreak, impact of infestation and Behaviour strategies for control. Sterile, self-sacrificing nymphs referred to as ‘Samurai’ aphids17, are seen amongst species of Hormaphidinae and Host range in this species also. They have tough and stout body, thic- kened forelegs and well-developed horns, giving them a C. lanigera largely feeds on plants of the family Poaceae. pseudoscorpion-like appearance. Certain percentage (20– There are a few records of the aphid colonizing plants 25) of first instar nymphs of C. lanigera attacked eggs belonging to Bixaceae and Combretaceae (Table 2). In and larvae of predaceous syrphid, Eupeodes and eggs of Japan, the species has been recorded on Miscanthus, but chrysopid, Chrysoperla pallens Rambur18. not on sugarcane10. C. lanigera secretes droplets containing alarm phero- mone from cornicles when attacked by the predator, Pseu- doscymnus kurohime (Miyatake). The number of aphids Life cycle and biology producing droplets differs based on the stage of the predator that attacks. When attacked by adults, 82% of C. The aphid undergoes an anholocyclic life cycle on Poaceae. 11 lanigera secreted droplets, whereas during larval attack, In Japan, Kurosu and Aoki found many alatae sexu- 19 only 7.2–12% secreted droplets . parae in the laboratory colony on sugarcane and stated that sexuparae played no significant role as they were nonfunctional. No host alternation has been reported in Seasonal occurrence Japan. Takahashi3 reported host alternating life cycle from Saccharum to Miscanthus by transfer experiments Population changes in relation to weather parameters, in Taiwan, even though no sexuals were noted in nature. time of planting and ratooning have been investigated for He studied the biology and found that the nymphal stage this pest. occupied ten days and alatae about 14 days. An early rainy season was the major cause for attack In Taiwan12, the optimum temperature for aphid deve- during the succeeding year in Indonesia20, and dry season lopment ranged from 20 to 23°C and aphids became inac- with high humidity was also conducive for aphid infesta- tive at temperatures below 15°C and above 28°C. The nym- tion21. Long periods of dry and wet seasons inhibited aphid phal developmental period varies under different tempe- population and alternating rainy and hot days enhanced rature and photoperiod regimes. Apterous nymphal stages population growth20,21. Severe damage occurred during occupied ten days3; 23–32 days12 in Japan and 15.8–16.5 high temperature and less precipitation in Taiwan22, days13 in Taiwan. Alatae, nymphal stage occupied 14 whereas dry season was favourable in the Philippines23.

Table 2. Host plants of C. lanigera recorded from India

Host plant Family Reference

Bambusa arundinaceae Retz. Poaceae 105 Bambusa sp. Poaceae 103, 106 Cynodon dactylon (L.) Pers. Poaceae 107 Grassum sp. Poaceae 108 Oplismenus sp. Poaceae 103 Saccharum officinarum Linnaeus Poaceae34,38–41 6, 101, 102, 105, 109–112 Themeda sp. Theaceae 113 Unidentified species Poaceae 112–114 Xylosma longifolia Bixaceae 113 Unidentified species Combretaceae 110, 115

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Table 3. Natural enemies of C. lanigera Zehntner Autumn-planted canes were more prone to heavy in- 24 Natural enemy Location References festation by the aphid in Taiwan , while in Japan it was 25 Parasitoids summer-planted canes that were prone to attack . Con- Hymenoptera Aphelinidae trary to the general observation of heavy infestation of Aphelinus desantisi Hayat India 35 sucking pests on ratoon crop, the woolly aphid was less Encarsia flavoscutellum Zehntner Java 39 Taiwan 41 abundant on ratooned sugarcane. The colonies started Netherlands 45 developing in November and December and then gradu- Philippines 116 Malaysia 93 ally increased during winter, with phenomenal increase India 35 Braconidae during summer. Diaeretiella rapae M’Intosh India 35 Attempts were also made in Taiwan to relate time of Diaeretus oregmae Gahan Philippines 36 Taiwan 37 planting with infestation levels by comparing the length Aphidius sp. Philippines 15 of aphid colony in spring- and autumn-planted crop26.

Encyrtidae The length of the colony developing on autumn-planted Anagyrus sp. India 35 Chalcididae cane was higher compared to spring-planted cane. Mea- Antocephalus sp. India 35 surements of the colony increased with decrease in tem- Predators perature from December to February. Coleoptera Coccinellidae Apterae were observed immediately after germination Anisolemnia dilatata (Fabricius) China 49 Cheilomenes sexmaculata (Fabricius) Philippines 15 and infestation increased until the minimum temperature India 77 reached 23°C in January-planted canes in Taiwan. The Coccinella septempunctata Linnaeus India 61 Coelophora biplagiata (Swartz) Vietnam 52 population peaked between the second half of April and China 117 the second half of June. Heavy precipitation coupled with Coelophora saucia Mulsant Formosa 12 Cryptogonus orbiculus (Gyllenhal) Philippines 48 high temperatures were the factors causing decline in the Curinus coeruleus Mulsant Philippines 118 Harmonia octomaculata (Fabricius) India 77 aphid population. In addition, typhoons before May were Pseudoscymnus kurohime (Miyatake) Japan 57 detrimental to the aphid27. Scymnus sp. Philippines 15 Synonycha grandis (Thunberg) Philippines 33 In Assam, the aphid incidence started in June and in- China 49 creased gradually to reach a maximum of 90.32% in Sep- Japan 50 28 Taiwan 14 tember and declined by January. In Uttar Pradesh, the Indo-China 52 29 Lepidoptera pest occurred during October to March . Lycaenidae The aphid emerged before sunset and continued to Taraka hamada (Druce) China 119 30 Pyralidae emerge until early morning . This probably helped aphids Dipha aphidivora (Meyrick) Malaysia 120 to escape desiccation due to high day temperatures. The China 59 India 35 spread of the aphid from one field to another thus took Taiwan 24 Japan 25 place mainly due to wind. Peak take-off occurred just Thiallela sp. Philippines 15 after sunrise, one day after emergence. Wind velocity of Diptera Syrphidae more than 6 cm/s increased the number of aphids alight- Dideopsis aegrota (Fabricius) Philippines 15 ing on the plant. India 77 Episyrphus balteatus (De Geer) India 77 Mature leaves were more prone to aphid attack com- Eupeodes confrater (Wiedemann) India 35 23 14 Eupeodes corollae (Fabricius) Philippines 15 pared to young leaves . Takano observed that narrow and Eupeodes kuroiwae (Matsumura) Japan 25 erect leaves were affected to a great extent while Patil5 Ischiodon scutellaris (Fabricius) India 77 Syrphus sp. Taiwan 24 found that soft, broad and drooping leaves were more Neuroptera suitable for aphid build-up. Chrysopidae Chrysopa sp. India 35 Cell-sap concentration was a major factor in inducing Chrysoperla carnea (Stephens) Philippines 33 31 Chrysoperla furcifera (Okamoto) Japan 92 susceptibility in different cane varieties . Varieties with Chrysopa pallens Rambur Philippines 33 low concentration of cell sap (averaging about 4.5° Brix) Japan 18 Dichochrysa alcestes (Banks) Japan 92 had lower infestation compared to those with high cell- Italochrysa acqualis (Walker) Japan 92 sap concentration (5.9° Brix). Susceptible varieties were

Hemerobiidae attacked less during rainy season due to drop in cell-sap Micromus sauteri Esben–Petersen Philippines 15 32 Taiwan 24 concentration, which was around 2 to 2.7° Brix . Irriga- Japan 25 ted fields suffered less infestation because of lower sap Micromus sp. India 78 Micromus timidus Hagen Japan 25 concentration. Healthy canes and fertile soil also contri- Hemiptera buted to the pest status. Anthocoridae Orius persequens (White) Java 78 Araneae Unidentified spiders Philippines 15 Nature of damage Pathogens Aspergillus sp. Philippines 15 Fusarium moniliforme Sheld. China 121 Initial aphid infestation was seen on the under surface of Penicillium oxallicum Currie & Thom. China 121 leaves along the midrib and then over the entire under

310 CURRENT SCIENCE, VOL. 87, NO. 3, 10 AUGUST 2004 REVIEW ARTICLES surface, covering it with flocculent, waxy secretion. Co- The parasitoids emerged in the early morning and their pious honeydew excretion often covers the entire upper longevity was reported to be 4–5 days. Females were surface of the leaves, leading to growth of sooty mould4. more abundant than males. Each female laid 1–6 eggs with Due to continuous sap-sucking, the crop becomes stun- an average of 2 in a single host. The egg, larval and pupal ted33, and continuous infestation leads to reduction in the stages lasted for 2–4, 5–10 and 6–11 days respectively41. length, circumference, weight and sugar content of the stalk, Hazelhoff42 developed a sampling method to determine and in susceptible varieties in Vietnam34, loss in tonnage per cent parasitism. He also emphasized the need for re- as well as sugar recovery35. Gupta and Goswami28 asses- distribution of the parasitoid from old to new plantations, sed the effect of 25 and 100% aphid-infested leaves on as the parasitoid is a poor flyer40. The system of crop rota- some yield and quality parameters of sugarcane and found tion is associated with the problem of forced migration of that 100% infestation had detrimental effects on the the parasitoid. Subsequent to the publication of notes on length (11.6% reduction), girth (3.5% reduction), weight utilization of the parasitoid in the field by Hazelhoff43,44, (16.6% reduction), length of internode (18.4% reduction) the parasitoid was evaluated in the field in the Nether- and width of leaf (4.9% reduction). Juice quality para- lands Indies45; however results of this trial are not known. meters also exhibited considerable reduction. The per cent The parasitoid was introduced into Taiwan46 and Phi- reduction in sucrose, brix, glucose, purity and commer- lippines23, but it failed to establish owing to the short cial cane sugar (CSS) was 53.3, 32.3, 25.3, 31.7 and 64.0 lifespan of the adult47 and heavy mortality of the adults respectively. Preliminary studies on loss estimation being during transit48. Though the parasitoid has been recorded conducted at University of Agricultural Sciences, Dhar- in India35, no studies have been made on its multiplica- wad indicate adverse effects of aphid infestation on yield tion and field evaluation. and quality parameters8. Predators Ant association Synonycha grandis (Thunberg) and Anisolemnia C. lanigera is attended by five species of ants and Poly- dilatata (Fabricius) rhachis dives Smith was the most abundant15. In addition to removing honeydew, the ant also protects the aphid from S. grandis was recorded as a potential predator of sugar- natural enemies. cane woolly aphid in Philippines33, China49, Japan50, Formosa51 and Indo-China52. S. grandis and A. dilatata Natural enemies have also been reported to feed on the bamboo aphid, Pseudoregma bambusicola Takahashi53. Seven species of parasitoids, 30 species of predators and Liu49 discussed the geographical distribution and rear- three species of fungal pathogens have been recorded on ing techniques of both the coccinellids that prey on C. C. lanigera (Table 3). Only in recent years, use of some lanigera. The optimum conditions for rearing were 26°C of these natural enemies for management of the pest has and 85% RH. The incubation, larval and pupal periods been attempted. averaged 4, 13.52 and 4.23 days respectively, for S. gran-

dis. The corresponding values for A. dilatata were 4, 13.16 and 5.16 respectively. Adult A. dilatata consumed Parasitoids on an average 87.8 aphids in an hour, the first instar larva, 100 aphids per day and grown up larva, 400–500 aphids Initial attempts to identify and exploit parasitoids were per day. S. grandis exhibited a little higher feeding poten- made as early as 1927. Encarsia flavoscutellum Zehntner tial. Oviposition took place several days after pairing in and Diaeretus oregmae Gahan were recorded from Java both coccinellids. S. grandis preferred to oviposit on both and Philippines respectively. D. oregmae was described surfaces of the leaf, and A. dilatata was found to lay eggs by Gahan36 in Ohio. Stary and Schlinger37 mentioned the on thin branches of bamboo. Pupation occurred on leaves. occurrence of the species from Taiwan. Among the two Pupae were the weakest stage in the life cycle and the parasitoids, E. flavoscutellum caused a natural parasitism prepupal stage was parasitized by an unknown chalcid49. of up to 30% in Java. Hazelhoff38 reported that there was S. grandis had 11 generations a year. The adult lived no need for adopting control measures as the parasitoid for 30–70 days in summer and 60–90 days in winter, and was prevalent even at early stage of aphid infestation. up to six months when the sexes were separated51. Pre- Techniques to enhance parasitization were outlined by mating period varied from 3 to 7 days, whereas preovi- Hazelhoff39, as he found that even during heavy aphid position period was 5–16 days. The oviposition period infestation, the parasitism reached a maximum of 20% lasted 1–2 months. The average fecundity was 673 eggs/ only. High humidity was necessary for effective multipli- female. Eggs were laid in masses of about 27. The opti- cation of the parasitoid40. mum temperature for egg-hatching ranged from 19 to

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22°C. Adult predators reared on agar solution and wheat aphid. However, the fourth instar larvae fed on all stages flour could not oviposit. Biological studies of this preda- of the aphid. The larvae attacked the nymphs from the tor were also conducted in Japan50 and India4 using C. underside of the head and thorax, whereas the adults attac- lanigera and C. silvestri respectively, as hosts. ked the abdomen of the prey57. However, no evaluation C. lanigera is most abundant on sugarcane from Octo- on its potential as a biocontrol agent has been made. ber to May, but the coccinellid appeared only after December, and was most abundant at the end of May51. Owing to difficulty in supplying sufficient aphids as Dipha aphidivora Meyrick food, attempts were made to rear coccinellids on artificial Surveys conducted in China from 1974 to 1984 indicated diets like 10% molasses, molasses with egg yolk, milk 54 D. aphidivora to be one of the most abundant and impor- powder and few drops of tomato juice , but these diets tant aphid-suppressing factors27. The predator has been failed. Coccinellids could be reared on aphids during recorded under three names, Isauria aphidivora, Cono- spring, though only 21% of the eggs hatched. In summer, bathra aphidivora and Cryptoblabes aphidivora from rearing on aphids was not possible perhaps because of high China, Malaysia and Philippines. Arakaki and Yoshiyasu58 temperature, when adult mortality was high and eggs failed 54 compared the morphology of the pyralid predators recor- to hatch . 2 ded on aphids to resolve the confusion in their identi- Field-released adults (at the rate of 500 per 200 m ) ten- fication. Crossing experiments and detailed studies on ded to migrate and only half of them were recovered 20 morphology indicated that all the species were conspecific days after release. The beetles disappeared two months and a new genus Dipha was proposed for this species. after release, without significantly reducing the aphid 54 The predator has also been recorded from Dimapur population . (Nagaland)35. Though it was recorded as a predator on Though the initial trials with S. grandis failed, research woolly aphid during 1978, serious efforts to multiply on its colonization and utilization was continued in China. 55 and use it for biological control were made only in 1997. Chen et al. mass-produced S. grandis on field-collected Chen et al.59 reared the pyralid at 25°C under laboratory woolly aphids and released them in the field. Takano and 51 conditions using freezed aphid nymphs stored at 1 and Noda suggested cold storage of adults and liberation at – 20°C. the outbreak centres at the right stage of aphid infestation. Deng et al.56 were able to store beetles at low temperature and release them at appropriate levels of infes- Pathogens tation. However, in these studies, released beetles could not bring about significant reduction in aphid population. Fusarium moniliforme Sheld. and Penicillium oxalicum Successful suppression of the woolly aphid by field Currie and Thom. have been recorded as fungal pathogens release of S. grandis was demonstrated by Deng et al.56 of C. lanigera in China60. Lingappa61 evaluated some after seven years of research. They collected S. grandis entomopathogenic fungi against the aphid and found and Coelophora biplagiata Swartz from sugarcane fields Metarhizium anisopliae to be effective in preliminary and stored them in the laboratory at temperatures ranging studies. Recent field trials conducted at Arabhavi, Karna- from 18 to 20°C and 75–85% RH. A total of 62.96 to taka with oil-in-water emulsion of B. bassiana and M. ani- 82.9% adult S. grandis survived after five months of sopliae caused 19.84 and 42.26% mycosis respectively62. storage. Survival rate was higher in C. biplagiata (45.7 to 100%). A total of 44,660 beetles were released into sugarcane fields, resulting in 90 to 96.6% reduction in aphid Defensive behaviour of predators population56. Aphids with aggressive behaviour are known in Horma- Species of Coelophora phidinae and Pemphiginae and 15 species of aphids have been recorded to be aggressive towards the predators62. C. biplagiata was introduced from Indo-China to Taiwan, C. lanigera and C. japonica (Takahashi) are known to where it established successfully. Later, it was taken from 52 attack immature stages of syrphids, chrysopids and coc- Taiwan and released in Java in 1925. Similarly, Coelo- cinellids18. phora saucia Mulsant was introduced to Taiwan from Predators have also evolved strategies to protect them- Tonkin and it established in the southern parts after ini- 12 selves from aggressive aphids. Larvae and adults of S. tial release . Information on its role in suppressing the grandis and A. dilatata are known to exude a toxic che- population of woolly aphid is not available. mical (reflex bleeding) from the femoro-tibial joints to protect themselves from soldiers of C. lanigera49. In addi- Pseudoscymnus kurohime (Miyatake) tion, both these predators frequently deposit their eggs The adults and first to third instar larvae of P. kurohime away from the aphid colony to prevent attack by the preferred to feed on the first instar nymphs of woolly aphid soldiers63.

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P. kurohime exhibits a typical egg-protection behavi- methods have been tried against sucking pests of sugar- our64. This predator lays eggs on the undersurface of leaves cane like Pyrilla perpusilla (Walker) and have proved near the leaf midrib in aphid colonies. The eggs are then successful74. Richards75 pointed out that stripping had covered with faeces-like secretion mixed with undigested other advantages like increase in sugar content as well as claws of the prey aphids. This behaviour not only pro- germination of seed sets. Gupta76, however, showed that tects the eggs from soldiers, but also protects it from high stripping has disadvantages, such as high cost and mecha- temperature and cannibalism. nical damage to eye buds by mammalian pests and field D. aphidivora spins a web around itself and feeds from workers, together with greater level of lodging. Thus, within this silken tunnel created for defence. The soldiers mechanical methods should be followed only after proper of C. lanigera get entangled in the web and fail to attack evaluation of the costs and benefits of the technique. the predator63.

Pest control strategies and future prospects Chemical control The predictions of potential distribution of the invasive A great deal of laboratory and field research on insecti- pests are helpful in developing strategies for monitoring cidal control of C. lanigera has been conducted in China, and managing serious pests. Recently, attempts have been Philippines and Taiwan. Lopez33 reported that spraying made to predict the spread of sugarcane woolly aphid. The with a contact pesticide, creoline was effective in control- results of these studies will be useful to fight the pest in ling the woolly. Sprays with 1% camphor oil, yam and bean an ecologically and economically sound manner9. oil, tea seed oil, tea decoction emulsified with sodium Deng et al.56 and Rabindra et al.77 advocated biological oleate and leaf extract of tobacco were also recommen- methods by augmentation of the coccinellids, S. grandis ded65. Tea seed oil was the most effective treatment, pro- and A. dilatata. The proposed methods are logical, as viding 93–99% mortality66. indiscriminate use of pesticides may lead to outbreak of Schradan, dematon67 and low-volume sprays with mala- secondary sucking pests. However, the methods need to thion68 were recommended in Taiwan and Philippines. Pan be justified economically, as mass multiplication techni- and Yang69 advocated aerial application of methyl deme- ques for both coccinellids are yet to be developed. Col- ton and dimethoate. Liu et al.70 also advocated application lection of predatory beetles from different locations and of dimethoate after June to control C. lanigera, without distribution where required is time-consuming as well as harming natural enemies. Fogging with a mixture of diaz- labour-intensive. In addition, predatory beetles are highly inon 60 EC and petrol at ratios of 2 : 3 and 1 : 2 was mobile. Earlier studies have indicated escape of predators effective, particularly when carried out on bright, wind- from the colonized areas in spite of availability of the less days in the afternoon or evening in Indonesia21. In host54. The other predator that is being proposed to be China, carbophos 40 EC resulted in more than 95% mor- utilized in India is the pyralid, D.? aphidivora77. Correct tality of C. lanigera71. identity of this predator needs to be established and For the control of recent outbreaks in Maharashtra and efforts should be made to mass multiply it in the labora- Karnataka, chemicals such as endosulfan, phosalone, mono- tory, conserve it in situ in the field and also redistribute it crotophos, dimethoate and metasystox sprays, and acephate to new areas of infestation. Micromus sp. is quite abun- and methyl parathion dust have been recommended5,61. dant and appears to be a potential candidate for mass multiplication and release78. This occurs even under low pest Host–plant resistance population feeding on first and second instars of aphid. Combination of Micromus sp. at low pest population and Attempts have been made to identify resistant sources in D.? aphidivora at higher population levels may work 16 sugarcane germplasm. Pan et al. studied the biology and well in the field. densities of aphid colonies on different varieties and iden- Parasitoids being more specific compared to predators, tified the variety ROC 1 to offer resistance to some extent. are better candidates as biocontrol agents of sedentary Similar efforts were made in Indonesia and a resistance species79 such as C. lanigera. Concerted efforts need to 72 breeding programme was started for C. lanigera . Two be made to explore the parasitoids of this species, such as hydroxamic acids, DIBOA and DIMBOA were detected in E. flavoscutellum in Northeast India; develop mass rear- sugarcane that may contribute to resistance to the aphid, ing techniques; assess their efficacy in the field and deve- 73 as they do to aphids on cereals . lop field-release and conservation strategies. The recommendations by Patil5 and Patil et al.80 in- Mechanical control clude proper sanitation, avoidance of transport of infested leaves from one area to another, removal of other alter- Patil5 advocated stripping of infested leaves and burning nate hosts in the vicinity, removal of infested leaves by them to avoid further spread. Such type of mechanical employing labourers, avoiding lodging of canes, burning

CURRENT SCIENCE, VOL. 87, NO. 3, 10 AUGUST 2004 313 REVIEW ARTICLES of trash after harvest and avoiding use of canes from in- M. and Uma Shaanker, R., Predicting the potential geographic dis- fested fields for sowing. These are general but important tribution of the sugarcane woolly aphid using GARP and DIVA- GIS. Curr. Sci., 2003, 85, 1526–1528. recommendations. The suggestion of ploughing after har- 10. Takahashi, R., On the aphids of Ceratovacuna in Japan. Kyontyß, vest seems baseless, as the pest neither overwinters nor lays 1958, 26, 187–190. eggs in the stubble. Recommendation of release of Chry- 11. Kurosu, U. and Aoki, S., Sexuparae of the sugarcane woolly aphid soperla carnea at the rate of 2500 larvae/hectare81 is not Ceratovacuna lanigera. Kyontyß, 1986, 54, 523–524. based on any systematic studies on the performance of the 12. Takano, S., On the biological control of sugarcane insects in For- mosa. Rep. Jpn. Assoc. Adv. Sci., 1941, 15, 231–233. predator against this pest in the field. C. carnea can be 80 13. Cheng, W. Y., Wang, Z. T. and Lin, K. J., Survival and reproduc- mass-produced easily and cheaply , but the predator has tion of woolly aphid on potted sugarcane. Report of the Taiwan 82 not been successful in suppressing C. lanigera and hence Sugar Research Institute, 2000, vol. 167, pp. 19–33. it is unlikely to contribute much to its management. 14. Takano, S., Relation of sugarcane varieties to insect pests. J. Sug- Sprays of the entomopathogenic fungus M. anisopliae arcane Plant. Assoc., 1937, 15, 195–198. 15. Rueda, L. M. and Calilung, V. J., Biological studies on the sugar- are being recommended and are also proving effective61,62. cane woolly aphid, Ceratovacuna lanigera Zehntner (Hemiptera: These recommendations make sense as many pathogenic Aphidoidea: Pemphigidae) on five varieties of sugarcane. 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