: FEEDINGPOTENTIALOF CASSIDA CIRCUMDATA HERBST(CHRYSOMELIDAE COLEOPTERA)ONIPOMOEAREPTANS(LINN.)(CONVOLVULACEAE )1 M. John George3 and Ipe M. Ipe2 (With one text-figure) Key words: Cassida circumdata Coleoptera, Ipomoea reptans, Convolvulaceae , ThetortoisebeetleCassidacircumdataHerbstfeedsontheaquaticplantIpomoeareptans(Linn.) (Family: Convolvulaceae), which grows profusely in the tropical wetlands. This plant species was found invading the open water area and changing the proportion ofhabitats available for aquaticbirdsand fishes. Thedifferentlarval instarsand adultsofthetortoisebeetlewerenoticed feedingontheleavesandtenderstemsofIpomoeareptans, oneoftheabundantaquaticplantsin Keoladeo National Park, Bharatpur, Rajasthan, which is among the majorprotected wetlands in India. Toassessthe impactofCassidacircumdataon itsmain hostplantI. reptans,consumption ofthe leaves in terms of area fed by different larval and adult stages were studied under laboratory conditions (30 ±2 °C and 50-70% relative humidity). The first to fifth instar larvae and adult differedwidelyintheirconsumptionofhostplantleaves.Theareaconsumedwasintheorder 1st <2nd<3rd<4th<5thinstars>adult. Significantcorrelationcouldbeobtainedbetweenthegrowth ofthe larvae and their food consumption. The different rates offood consumption indicate the varying energy requirements ofthe various larval stages ofthe beetle. Introduction Keoladeo National Park of which the most dominantspeciesisagrass,Paspalumdistichum , Recently, much attention has been given that covers amajorpart ofthe aquatic area. The to the herbivore-plant relationship, with restis mostly covered withIpomoea reptans an , reference to its ecological and evolutionary important amphibious herb. Ipomoea reptans impacts (Denno and McClure 1983). Such floats inwater, usuallyappears as atrailingherb studies demand a knowledge of the actual and changes the proportion of the habitat consumption of the host plant parts by the available for aquatic birds and fishes (Ali and herbivore and its growth rate. The quantitative Vijayan 1986). In the Park, I. reptans remains assessment ofconsumption ofhostplants by its in a dormant stage during winter (December- pestmayhelp us to assess the impact ofthepest February). on the growth of the plant and thereby the Insummer(April-June), as thewaterlevel possibleuseofthatpestasanagentofbiological drops and the stem makes contact with the control. ground, it produces leaves in large numbers. The tortoise beetle Cassida circumdata Aftermonsoonandwhenthelevelofwaterrises, Herbst feeds on the aquatic plant Ipomoea I. reptans spreads andattains maximumgrowth. reptans (Linn.), which grows profusely in Thispaperreports the resultofthe study carried tropical wetlands. About 91 species ofwetland outin the field andunder laboratory conditions, macrophytes have been identified from the to determine the feeding potential of 'AcceptedJuly,1999 C. circumdata. 2SchoolofEntomology,St.John’sCollege, The life-cycle of C. circumdata includes Agra282002,UttarPradesh,India. 5 larval stages and apupal stage. The eggs were ?PPerreusmenbtavAododrre,sEs:maMkaurlTahmo6m8a3C5o4l2l,egKeerfaolraW,oImndeina., laidontheventralsideofI. reptansleaves.Under 370 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 97(3), DEC. 2000 FEEDINGPOTENTIALOFCASSIDACIRCUMDATA laboratory conditions (30 ±2 °C), the eggs food consumed by the adult and grub ofLema hatched within 4 days. The duration of the 5 lacordairei (Chrysomelidae), Visalakshi and larval instars was 2, 1-2, 2, 2 and 4 days Nair (1987) used similar methods. respectively. The pupal stage lasted for4 days. Alongwiththeexperimental studies inthe laboratory, an attemptwas made to quantify the Study area feeding impact ofdifferent instars ofthe beetle on the host plant. This was carried out by The present study was carried out in the collecting the plant and different stages of the well-known protected wetland, the Keoladeo beetles fromrandomlyplacedquadrats of100 x National Park, Bharatpur (27° 7.6' to 27° 12.2' 100 sq. cm. Collectionsweremade from6 study N and77°29.5' to 77°33.9' E), Rajasthan, India. plots. Oncethequadratwasplaced, alltheleaves above water were plucked and transferred into Methodology plastic bags (harvest method). The adults that flew offon being disturbed were also counted. Both sexes were almost alike, the male In the laboratory, all the leaves were carefully being slightly smaller than the female. To study examined and the different instars and adults the feeding and oviposition, experiments were counted. The leafarea was measured using the earned out in plastic containers (height = 8 cm same method as above. The leaf biomass was anddiameter=3 cm)withscrewtophavingholes quantified using an electrical monopan balance forventilation. Discs offilterpaperwereplaced of sensitivity 0.1 mg. The samples were taken at the bottom ofthe containers to absorb excess once in seven days and monthly averages were moisture and facilitate cleaning. Mating pairs calculated. were collected from the field and kept in these containers. The leaf-area fed by the adult and Resultsand discussion the eggs laid after 24 hrs were recorded. The adults, when kept in containers laid eggs on the Labexperiment: The average leafarea of leavesofI. reptans, andtheeggshatchedwithin Ipomoea reptans consumed and the average 4 days. growth (length and breadth) of the larvae of Newly emerged larvae wereplacedone in Cassidacircumdataforeachdayaresummarized eachcontainerandprovidedwithfreshleaves of in Table 1. The maximum leaf area was I. reptans daily. The body length and breadth consumed on the eighth day by the 5th instar were measured every 24 hrs using a compound larva (59.33 sq. mm). The maximum increase microscopewithacalibratedocularmicrometer. in the larval stage was noted on the seventh and When fresh leaves were freely available, the eighth day. On the ninth day, the larvae slowly larvae and adults fed upon the fresh area. The decreased their rate of feeding and entered leafareaconsumedandremovedatonemeal(leaf pupation. The 5th instar larva fed about mm area fed at a stretch) was measured by placing 101.16 sq. leaf area within 3 days. The the leafon graphpaper and tracing the leafarea maximumarea was feduponby the two day old fed, and then counting the squares of the leaf 5th instar larva. This study showed that the 5th area fed, as suggested by Simmonds (1949). As instar larvae can consume almost 4 times more theearlyinstars fedvery little andmade circular than the 4th instar and about 100 times more holes, the radius of the circular path fed was than the 1st instar (Table 1). measured with graph paper and by a calibrated, A single larva from the 1st through the mm ocular micrometer. To estimate the quantity of 5th instar consumed about 145.95 sq. leaf JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 97(3), DEC. 2000 371 FEEDINGPOTENTIALOFCASSIDACIRCUMDATA Table 1 FOODCONSUMPTIONANDGROWTHOFTHELARVAEOFC. CIRCUMDATAREAREDON/.REPTANS INTHELABORATORY(30±2°C) Days 1 2 3 4 5 6 7 8 9 10 11 Stage(instar) 1st 2nd 3rd 3rd 4th 4th 5th 5th 5th Pupa Pupa Increaseinlength(mm) 0.15 0.28 0.3 0.2 0.45 0.45 0.75 0.58 0.18 -0.08 0 Increaseinbreadth(mm) 0.08 0.17 0.15 0.13 0.37 0.17 0.38 0.55 0 0 0 Areafed(sq.mm) 1.86 5.01 4.68 9.83 9.65 16.79 30.25 59.33 11.58 0 0 area ofI. reptans within 9 days (Table 1). The Table2 average leafarea collected during July-October LEAFTAHREELAAORFVAI.EREAPNTDAANDSUCLOTNSC.UCMIERDCU(MSQD.AMTMA)BY was 17.55 sq. cm. The study showed that two AT30±2°C(N=80) larvae from the 1st instar through 5th instar can completely eat and skeletonise a single leaf. 1stinstar 0.21 ±0.13 To study the feeding behaviour of the 2ndinstar 0.35 ±0.19 3rdinstar 0.80 ±0.37 beetle, thehostplantleaveswereobservedwhen 4thinstar 1.86 ±0.82 there was less attack. Ofthe total holes fed and 5thinstar 5.32 ±3.41 made by the larvae and the adult (N 1 5,265 Adult 3.77 ±1.72 holes) 95.5% of the holes indicated only one feeding. This indicatesthatthe larvae andadults was noted on the 16th day. In Aspidomorpha preferredtofeedonfreshareawhenthehostplant miliaris, the adult consumed seven times more leaves were plentiful. leaf area than the 5th instar (Manjunatha et al. The leafareafedandremovedatonemeal 1987). BombyxmoriandProtoparcesexta, both (in a stretch) by each instar larva and adult are lepidopterousleaffeeders,eatabout97%oftheir summarized in Table 2. The result showed that total intake during the lasttwo instars andabout the 5th instar larvae consumed greater leafarea 99% duringthe lastthree instarsrespectively. In at a single meal and the first instar the least. Bombyx mori it was noticed that the efficiency , The adult C. circumdata pair under ofstorageofmetabolizableenergyincreaseswith mm captivity (N=15) fed on 56.81 ±12.53 sq. age, reachingitspeakinthe 5thinstar. The large leaves and laid 12.6 ±6.1 eggs within 24 hrs. amount ofenergy stored during 5th instar is, of The mating pairs fed 17.1 ±4.8 times within course, needed to support the non-feeding pupa 24 hrs. The area fed upon by the mating pairs and adult (Waldbauer, 1968). waslesscomparedtothe5thinstarlarva, asmore Field experiments: Field observations time was utilized for mating and laying eggs. revealedthatthebeetle appears inthe study area No significant correlation was noticed between during March (mean min. temp. = 19 °C). The the area fed and the number ofeggs laid. host plant was available only along the dykes. Simmonds (1949) assessed the leaf area The population of C. circumdata remained consumed each day by the larvae ofPhysonota almostconstantduringthepeaksummermonths. alutacea(Cassidinae).Theleafareaconsumption With the onset of monsoon and the release of wasseentoincreasewiththegrowthofthelarvae water from Ajanbund, an inundation reservoir eachday, themaximumconsumptionofleafarea situated0.5kmsouthofthePark,theplantspread 372 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 97(3), DEC 2000 FEEDINGPOTENTIALOFCASSIDACIRCUMDATA over the area by vegetative reproduction. There be observed due to low population ofthe beetle is spatialvariation inthe abundance ofthisplant (George 1988). The maximumpopulation (468/ in the Park. By this time, the beetle had settled sq. m) and leafarea (17.8 sq. mm) during 1986, indifferentareas.Theadultsmainlyconcentrated was noticed in September. in certain areas, fed and laid eggs. There was Kolodny-Hirsch and Harrison (1982) wide fluctuation in the population of different conductedfieldandfieldcagestudiestocompare stages of the beetle among the different study larvalinjurybythetobaccobudworm,Heliothis plots. Afterlayingeggs, the adultsmovetoother virescens and corn ear worm, Heliothis zea less infected areas. The adult beetles prefer to (Lepidoptera: Noctuidae). Their observations lay eggs on the underside offreshyoung leaves. showedthattheplantcompensatedforleaflessby Eggs were never laid on skeletonised leaves, to increasingthelaminalareaofthedamagedleaves. ensure enough food for the emerging larva. The population of C. circumdata in the , Theaveragemaximumnumberofdifferent Park,reachedapeakinSeptember,andasaresult stages ofthebeetlewasnotedduring September the leafarea and biomass declined. In the end, (Fig. 1). Theleafareaofthehostplantwasfound onlyleafskeletonsremainedintheinfestedareas. following the same trend as that ofthe beetle. Thus, leaf biomass became zero towards the Thus, a significant positive correlation was onset ofwinter. In winter, both the plant and its obtained between biomass and leaf area ofthe pest remained dormant. The beetles were found plantwiththepopulationofthebeetle(r=0.821, hibernating, in the Park, on terrestrial plants, p - 0.001; r - 0.8472, p = 0.0001, n= 60). namely Salvadora persica, albeit in small It was observed that the host plant numbers(George 1988).Thisformedthebreeding compensated for the leaf area fed upon by the stock for the next population during February- larval and adult C. circumdata by producing March, and by this time the hostplant also starts larger leaves. Leaves with greater surface area producing smaller leaves. Thus, the interrelated were noticed in August and September during life-cycles ofthehostandpest continue. 1985, when the population ofthe beetle was at The study unravels a unique relationship itspeak(Fig. 1). Butinalaterstudyduring 1986- between the plant and the beetle. Though the 87, asimilarresponse ofthehostplantcouldnot beetle controls the biomass ofthe host plant, it POPULATION/AREA 10000 § LEAFAREA TOTAL 1000 POPULATION 100 = Fig. 1: Leafarea(sq. mm) ofI. reptans and population ofC. circumdata JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 97(3), DEC. 2000 373 FEEDINGPOTENTIALOFCASSIDACIRCUMDATA does not completely destroy them, thereby sponsored by the US Fish and Wildlife Service ensuring its own survival. The most important through the Ministry of Environment and barrierswhichpreventthebeetlefromdestroying Forests,Govt,ofIndia. Encouragementreceived the plant entirely are the ability ofthe plant to fromMr.J.C.Daniel,Hon. Secretary,BNHSand survive under water and low ambient tempera- Dr. V.S.Vijayan,Director,SACON,Coimbatore tures (average minimum temp, in December- is acknowledged. We also thank Dr. N.K. February 8 °C). Ramachandran, Wildlife Institute of India, for criticism and suggestions, and Dr. C.R. Acknowledgements Ajithkumar, PrincipalInvestigator, Mr. K. Raju Thomas and Mr. C.R. Biju, Research Scholars, Thispaper is an outcome ofthe Keoladeo BNHS Research Station, Kalady, Kerala, for National Park Ecology Project of the BNHS, their help. Refer • N CES Ali, S. &V.S. Vijayan(1986): KeoladeoNational Park Manjunatha,M.,G.T.T.Raju,D.N.R.Reddy&S.K.Dutta EcologicalStudySummaryReport,1980-85.Bombay (1987): Studies on the biology ofAspidomorpha NaturalHistorySociety,Bombay,pp54. miliarisF.(Cassididae:Coleoptera)onthreespecies Denno,R.F.&M.S.McClure(1983):Variableplantsand ofIpomoea and leafarea consumption studies on Herbivores in Natural and Management Systems. IpomoeaangulataLamk.J.Bombaynat. Hist. Soc. AcademicPress,Inc.,London,pp717. 84(3):704-708. George,M.J.(1988):Bioecologyandpopulationdynamics Simmonds, F.J. (1949): Insects attacking Cordia of Tortoise beetle Cassida circumdata Herbst macrostachya(Jacq)Roem.&Schult. in theWest (Chrysomelidae:Cassidinae)inKeoladeoNational Indies, I. Physonota alutacea Boh (Coleoptera : UPnairvke,rBshiatyr,atApgurra..Ph.D.ThesissubmittedtotheAgra VisalaCakssshiid,inAa.e&).MR.epRr..G.fKr.omNCaainr.(E1n9t8o7)m:olB.ioVlolo.gy8o1f,NLoe.m8a. Kolodny-Hirch, D.M. & F.P. Harrison (1982): lacordairei Baly. (Coleoptera : Chrysomelidae : Comparativedamageandleafareaconsumptionby Criocerinae)apestofYamDioscoreaalatainKerala. the Tobacco bud worm and Corn ear worm on Entomon. 3(1): 129-131. Maryland Tobacco. Econ. Entomol. 75(2): 168- Waldbauer,G.P.(1968):Theconsumptionandutilisation 196. offoodbyinsects.Adv. Insect.Physiol. 5:229-288. 374 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 97(3), DEC. 2000