JAE 012 "0888#

J[ Appl[ Ent[ 012\ 236Ð249 "0888#Þ 0888\ Blackwell Wissenschafts!Verlag\ BerlinISSN 9820!1937

Suitability of physic nut (Jatropha curcas L.) as single host plantfor the leaf-footed bug Leptoglossus zonatus Dallas (Het.,Coreidae)C[ Grimm and A[ Somarriba

Institute of Forest Entomology\ Forest Pathology and Forest Protection\ University of Agricultural Sciences\Vienna\ Austria

Abstract] The leaf!footed bug Leptoglossus zonatus "Dallas# "Het[\ Coreidae# was reared in Nicaragua on a diet consistingof unripe physic nut "Jatropha curcas L[\ Euphorbiaceae# fruit only[ The suitability of this host plant is described by themortalities in the developmental stages from egg to adult\ which are presented in a life table\ and the durations of thedi}erent stages[ Pre!oviposition time\ number of ovipositions and total number of eggs per female are given[ Biometricmeasurements of all instars were made[ Protein\ oil and water content of fresh physic nut fruit were measured throughouta whole year[ The results show that physic nut is a highly suitable host plant\ which can maintain populations of thisinsect pest throughout the whole year as fruit are constantly present with only little ~uctuation in their nutritionalquality[

0 Introduction

The leaf!footed bug Leptoglossus zonatus is a commonspecies in North and South America\ with a geo!graphical range that extends from the south!westernUnited States to Argentina[ It is a pest of a wide rangeof host plants\ including the basic food crops maize andsorghum\ many vegetables like tomato and all kinds ofcucurbitaceas\ and fruit including citrus\ passion fruit\pomegranate and cashew nut "GRIMM and MAES\0886#[ In past years physic nut "Jatropha curcas# plan!tations have been established in Nicaragua as a sourceof biofuel\ which is produced at an industrial level bytransesteri_cation of the seed oil obtained from thisperennial tree crop "FOIDL et al[\ 0885#[ One of the mostfrequent pest species found in physic nut plantations isL[ zonatus[ Ovipositions\ which are laid in chains of upto 49 eggs\ are found on stems\ branches\ leaves andpeduncles of physic nut[ Both the gregarious larvae andthe adults have been observed to feed on leaves andunripe fruit\ provoking premature abortion and thedevelopment of malformed and hollow seeds[ Eggs andnymphs\ however\ are also found on many other cul!tivated and wild herbaceous and woody plants in andaround physic nut plantations\ and the higher frequencyof adults compared with eggs and nymphs on physic nuttrees indicate an immigration from other hosts "GRIMM

and FUÝHRER\ 0887#[The physic nut tree is well known for its insecticidal

qualities\ which a}ect insect species from a wide rangeof families[ Ingestion with the diet has been found toinhibit growth of lepidopterous pests "SAUERWEIN etal[\ 0882^ MENGUAL\ 0886#[ Contact toxicity andchemosterilizing e}ects were described by SOLSOLOY

"0882# and SOLSOLOY and SOLSOLOY "0886# on Homo!

U[ S[ Copyright Clearance Center Code Statement] 9820Ð1937:88:1295Ð9236 , 03[99:9

ptera\ Coleoptera\ Lepidoptera and Diptera[ PATIL andSINGH "0880# report the traditional use of J[ curcas asa fumigant against bed bugs "Cimex lectularius L[\ Het[\Cimicidae# in Ghana[ The insecticidal agents are con!centrated in the oil\ which suggests that the physic nutfruit may not be an ideal host for polyphagous bugs thathave not evolved a speci_c resistance to the chemicalcompounds that constitute the pesticidal mode ofaction[

To improve management strategies of heteropterouspests in physic nut plantations it became necessary todetermine whether L[ zonatus can complete its entirelife cycle on a diet consisting solely of physic nuts orwhether other host plants are necessary[ To furtherestimate the suitability of physic nut\ the biometric andbionomic data of insects reared on physic nut werecompared with data obtained by other authors ondi}erent host plants[

1 Materials and methods

Adults of L[ zonatus were obtained from physic nut plan!tations in Nicaragua and fed with a diet of freshly picked\unripe physic nut fruit\ that were replaced on alternate days[No additional diet or water was given[ The insects were keptin wooden cages "29 cm × 29 cm × 39 cm# with plastic gauzewalls\ that were placed in an open\ roofed space\ which leddirectly to an experimental plantation\ so that the climaticconditions were similar to those found in the adjoining physicnut stands[ Daily means of relative humidity in the rearingfacility ~uctuated between a minimum of 58) and amaximum of 67) and temperatures varied between 15[6 and20[0>C[

The quality of the diet was monitored by taking randomsamples of 19 fruit per week throughout the year beginning

237 C[ Grimm and A[ Somarriba

Table 0[ Life table of _ve ovipositions of Leptoglossuszonatus

Developmentalstage Lx Ix Dx dx qx

******************************Eggs 108 099[9 54 18[6 18[6First instar 043 69[2 37 10[8 20[1Second instar 095 37[3 27 06[3 24[7Third instar 57 20[0 1 9[8 1[8Fourth instar 55 29[0 1 9[8 2[9Fifth instar 53 18[1 1 9[8 2[0Adults 51 17[2 51 17[2 099[9

Lx\ number of surviving at beginning of stage^ lx\ survival atthe beginning of stage per 099 eggs^ Dx\ absolute mortality^ dx\mortality per 099 eggs^ qx\ mortality per 099 at beginning of stage[

in July 0884[ All fruit were collected in plantations in andaround Managua\ Nicaragua[ The dry weight of the fruitwas measured with semi!analytical scales "Sartorius L1199S\Go�ttingen\ Germany#[ The water content was measured byweighing and drying at 79>C for 37 h[ Oil and protein contentof the seeds were analysed according to the proceduresdescribed by REITINGER "0882#[

Five ovipositions from females kept in the rearing facilitiesdescribed above were placed into individual cages[ The num!ber and stage of nymphs were recorded daily until the _nalmoult[ From these data life tables and the duration of eachstage were computed[

Five pairs\ each consisting of a male and female adult fromdi}erent parents\ were also placed in individual cages afterthe _nal moult[ Pre!oviposition time and survival time wererecorded as well as the number of ovipositions per female andthe number of eggs per laying[

At each developmental stage\ 49 individuals were collectedand subjected to measurements with a stereo microscope"Wild 2MC\ Heelbrugg\ Czech Republic# and the cor!responding Leica graticule[ These included the total width ofthe head capsule\ the width between the eyes\ the maximumlength and width of the pronotum and overall length of theinsects[

2 Results

The dry weight of the fruit oscillated between a mini!mum of 0[92 g in the dry season and a maximum of1[90 g in the rainy season with a mean of 0[65 g"SD�9[322#[ The water content lay between 71 and80) "mean � 74[70)^ SD�1[193#[ The protein con!tent of the seeds was very constant\ ranging from 02[72to 03[53) "mean�03[29)^ SD�1[284#[ The oil con!tent of the seeds showed the highest ~uctuations of allmeasured parameters[ The minimum found was 5[75)and the maximum 16[80) "mean�04[71)^SD�5[687#[

The percentage hatch of eggs was 69[2) and the totalmortality of nymphs was 48[6)[ Second instars had thehighest mortality at 24[7) "table 0#[ The eggs took anaverage of 7[3 days to hatch[ Total larval developmenttook 14[5 days\ in which the _rst stage was shortest withan average of 1[4 days and the _fth stage was longestwith 5[7 days "table 1#[

Mean survival time was 72[5 days for male adults and

Table 1[ Development time in days of all stages of Lep!toglossus zonatus

Stage Mean SD min max n******************************

Eggs 7[33 0[905 6 01 043First instar 1[37 9[462 1 3 095Second instar 4[96 1[909 2 02 57Third instar 3[35 0[068 1 6 55Fourth instar 3[31 0[146 1 6 53Fifth instar 5[72 9[860 3 8 51Total larval 14[51 3[086 19 21[4 51

developmentMale adults 72[5 37[230 21 037 4Female adults 76[3 11[677 52 003 4Pre!ovipostion time 13[7 5[232 19 24 4

SD\ standard deviation^ min\ minimum^ max\ maximum^ n\ num!ber of observations[

Table 2[ Oviposition data for Leptoglossus zonatus

Stage Mean SD min max n******************************

Ovipositions 6 0[114 5 8 4Eggs per oviposition 21[6 01[794 01 46 24Eggs per female 118 84[496 033 268 4

SD\ standard deviation^ min\ minimum^ max\ maximum^ n\ num!ber of observations[

76[3 days for female adults[ The pre!oviposition timewas 13[7 days "table 1#[ Females laid an average of 118eggs in seven ovipositions "table 2#[

The biometric measurements of the di}erent instars\male and female adults are given in table 3[ Femalesare larger than the males[ The gender di}erences of allmeasured parameters are signi_cant at the P³ 9[990level according to Student|s t!test[

3 Discussion

The data presented show that L[ zonatus can live andreproduce on a diet consisting solely of unripe physicnut fruit with no access to free water[ Despite its proveninsecticidal qualities\ physic nut is as good a food sourceas other known host plants\ which can be shown bycomparing the survival rates\ developmental time andsize of the di}erent instars with data obtained by otherauthors using maize\ soybean\ green beans and sorghumas diet[

PANIZZI "0878# reared L[ zonatus on diets of greenmaize\ soybean and bean pods at a temperature of142 0>C\ 542 4) relative humidity and a 03 h pho!toperiod[ Mortality of the second to _fth instars rangedfrom 49[9) with maize to 74[9) on bean pods\ com!pared with 30[4) obtained in the experiments on physicnut[ Total development time from second instar to _nalmoult was shortest on maize with 31[9 and 32[3 daysfor males and females\ respectively[ The correspondingvalue for physic nut is 19[7 days[ The mean number of

238Physic nut as a single host plant for the leaf!footed bug

Table 3[ Head capsule width\ inner eye distance\ lengthand width of pronotum and total length of different stagesof Leptoglossus zonatus[ All data in millimetres

Mean SD min max n******************************

First instarsHead capsule width 9[61 9[956 9[49 9[78 49Distance between eyes 9[34 9[927 9[27 9[51 49Width of pronotum 9[63 9[955 9[42 9[73 49Length of pronotum 9[10 9[951 9[01 9[32 49Total length 1[67 9[347 1[93 2[69 49

Second instarsHead capsule width 9[88 9[949 9[80 0[07 49Distance between eyes 9[59 9[947 9[37 9[63 49Width of pronotum 9[74 9[956 9[63 0[95 49Length of pronotum 9[28 9[945 9[15 9[49 49Total length 4[96 9[477 2[73 5[95 49

Third instarsHead capsule width 0[15 9[983 0[93 0[56 49Distance between eyes 9[63 9[960 9[37 9[82 49Width of pronotum 0[06 9[039 9[63 0[56 49Length of pronotum 9[51 9[986 9[26 9[60 49Total length 6[00 9[871 3[44 8[28 49

Fourth instarsHead capsule width 0[55 9[963 0[41 0[70 49Distance between eyes 9[86 9[958 9[63 0[00 49Width of pronotum 1[90 9[068 0[37 1[30 49Length of pronotum 0[96 9[019 9[63 0[15 49Total length 00[14 0[932 8[27 02[93 49

Fifth instarsHead capsule width 1[19 9[093 0[77 1[25 49Distance between eyes 0[18 9[972 0[93 0[30 49Width of pronotum 3[07 9[303 2[00 3[71 49Length of pronotum 1[19 9[157 0[42 1[51 49Total length 04[68 0[289 02[93 08[97 49

Male adultsHead capsule width 1[30 9[090 1[93 1[44 49Distance between eyes 0[20 9[970 0[00 0[33 49Width of pronotum 4[58 9[252 4[94 5[35 49Length of pronotum 2[66 9[244 2[92 3[44 49Total length 07[23 9[711 05[69 19[56 49

Female adultsHead capsule width 1[48 9[982 1[26 1[67 49Distance between eyes 0[34 9[968 0[07 0[52 49Width of pronotum 5[33 9[272 4[45 6[26 49Length of pronotum 3[02 9[248 2[12 3[64 49Total length 19[69 0[156 06[70 12[95 49

SD\ standard deviation^ min\ minimum^ max\ maximum^ n\ num!ber of observations[

eggs laid per female fed on a maize diet was 096[5 in4[1 ovipositions^ on soybean only one oviposition wasobtained[ With the physic nut diet\ on the other hand\ anaverage of 118 eggs were obtained in seven ovipositions[Percentage egg hatch was 64[7) on maize and zero onsoybean\ whereas 69[2) of the eggs obtained with thephysic nut diet hatched[ Pre!oviposition time in maizewas 11[9 days compared to 13[7 days on physic nut[

The development of L[ zonatus on diets of maize andsorghum was studied by MATRANGOLO and WAQUIL

"0883# at 172 1>C\ 672 4) relative humidity and a01 h photophase[ Total nymphal development took 17[6days on maize and 20[5 days on sorghum comparedwith the 14[5 days observed on physic nut[ Mortality of

Fig[ Comparison of overall lengths of different instarsreared on diets of physic nut and maize "FERNANDES

and GRAZIA\ 0881#[ Error bars indicate 2 0 standarddeviation

nymphs totalled 42[7) on a maize diet and 44[0) whenfed on sorghum[ Mortality on physic nut was 48[6) forthis developmental period[ Adults were fed on sorghumonly by MATRANGOLO and WAQUIL "0883#^ femalelongevity was 60[9 days and males lived an average of43[2 days[ On physic nut the data were 76[3 and72[5 days\ respectively[ Pre!oviposition time of femalesfed with sorghum was 31[5 days\ which is longer thanthe 13[7 days observed with the physic nut diet[ Thenumber of ovipositions obtained per female was 4[4 andthe total number of eggs 85[1 compared to seven and118 on physic nut[

JACKSON et al[ "0884# reared L[ zonatus on a greenbean diet at 39 to 59) relative humidity\ with a pho!toperiod of 05 h light[ At 29>C the nymphal mortalityamounted to 69[6)\ which is higher than that observedin the studies with physic nut\ and total nymphal devel!opment time to adult was 33[5 days\ which is longerthan on physic nut under natural conditions[ Adultslived for an average of 68[3 days\ which is shorter thanthe life span obtained for either males or females in thisstudy[

Measurements of the nymphs of L[ zonatus rearedon maize were published by FERNANDES and GRAZIA

"0881#[ Overall length was larger for the insects rearedon physic nut up to the fourth instar "see _g[#\ and thehead capsule was wider for the insects reared on physicnut up to the third instar[ The distance between the eyeswas larger for the insects reared on maize in all instars^The width of the pronotum was wider for third andhigher instars reared on maize\ but the length of thepronotum was larger for fourth and _fth instars rearedon physic nut[

The climatic conditions prevalent in plantations inthe Paci_c lowlands of Nicaragua are near the optimumfor the development of this species[ Annual averagetemperature in Managua is 15[4>C\ with monthly meansranging from 14[1>C in December to 17[5>C in April

249 C[ Grimm and A[ Somarriba

"FENZL\ 0877#[ JACKSON et al[ "0884# found anoptimum temperature for the development and repro!duction of L[ zonatus between 14 and 29>C\ and thehighest percentage of hatched eggs between 04 and19>C[ No nymphs reached adulthood at a temperatureof 04>C[

Physic nut plantations in Nicaragua provide an excel!lent environment for the build!up of populations of L[zonatus[ All measured parameters show that physic nutis equally or better suited than any of the other dietspreviously used to rear this bug[ Furthermore\ physicnut fruit are found in plantations in and around Man!agua throughout the whole year with very little ~uc!tuation of its nutrient contents\ particularly protein[Even though immigration from other wild and cul!tivated host plants has been observed\ the control ofL[ zonatus cannot rely on elimination or treatment ofalternative hosts alone but must include control activi!ties in the plantations themselves[

Acknowledgements

Research was conducted at the Proyecto Biomasa in Nica!ragua\ a development aid project executed by Sucher andHolzer Ges[m[b[H[ on behalf of the Austrian Ministry ofForeign A}airs[

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Authors| addresses] Dr CHRISTOPH GRIMM "correspondingauthor#\ Novartis Crop Protection AG\ CH!3991 Basel\ Swit!zerland^ Ing[ ANTHONYETTA SOMARRIBA\ Departamento deBiomasa\ Universidad Nacional de Ingenier(�a\ AP 321\Managua\ Nicaragua