Occurrence of Fruit Flies of the Genera Anastrepha and Ceratitis (Diptera: Tephritidae), and Their Host Plant Availability

in Costa Rica Author(s): Luis Fernando Jiron and Ingemar Hedstrm Source: The Florida Entomologist, Vol. 71, No. 1 (Mar., 1988), pp. 62-73 Published by: Florida Entomological Society Stable URL: http://www.jstor.org/stable/3494894 . Accessed: 25/06/2013 15:43
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62

Florida Entomologist 71(1)

March, 1988

OCCURRENCE OF FRUIT FLIES OF THE GENERA ANASTREPHA AND CERATITIS (DIPTERA: TEPHRITIDAE), AND THEIR HOST PLANT AVAILABILITY IN COSTARICA.
Luis FERNANDOJIRON

Escuela de Fitotecnia, Museode Insectos, Universidadde Costa Rica, CiudadUniversitariaRodrigoFacio, San Jose, Costa Rica
AND

INGEMARHEDSTROM

Departmentof Zoology, Section of Entomology, Uppsala University, P.O. Box 561, S-75122 Uppsala, Sweden and Escuela de Biologia, Universidadde Costa Rica, CiudadUniversitariaRodrigoFacio, San Jose, Costa Rica
ABSTRACT

Collectionsof wild and cultivated tropical fruits were examinedfor infestationby Anastrepha spp. and Ceratitis capitata (Wiedemann)(Diptera:Tephritidae).Of 440 collections(with at least 4 fruits in each)from201 sites throughoutCosta Rica, Central America,95%of the fruit flies (n = 4126)belongedto the generaAnastrepha(A. obliqua, A. striata, A. serpentina, A. manihoti, A. pickeli, A. distincta, A. chiclayae, and A. fraterculus), while 4.7%were C.capitata.Anastrephaspp. had a strong preferencefor host plants within the same family.A. obliquainfested 92%of the attackedMangifera indica (mango),87.5%of the Spondias mombin(jobo),and 100% of the S. dulcis (June A. plum/goldenplum) and S. purpurea (Spanishplum) collections,all Anacardiaceae. striata was recovered from 97.8%of the infested Psidium guajava (commonguava), 97% of the P. friedrichsthalianum (Costa Rican sour guava) and 100%of the P. savanarum (guiisaro) collections;these three host species belong to the Myrtaceae.A. serpentina was the dominantspecies in Sapotaceaehost plants, and it was recovered from 100%of all infested Manilkara achras (zapote), and Pouteria cainito (yellow caimito), and 98% of Chrysophyllumcainito (caimitomorado).A. manihoti and A. and 54.2%infestation, pickeli were associatedwith Manihotesculenta (cassava)(45.8% respectively). A. chiclayae infested 100%of the Passiflora quadrangularis(granada) collections.A. distincta infested species of Inga (Fabaceae)exclusively.C. capitatawas the only species of fruit fly recoveredfromPrunus persica (clingpeach).The infestation rates for C. capitata were low (<7%). Annualphenologyof the abovelisted host plants is presented with additionalobservationson interactionsbetween fruit flies and their host plants.
RESUMEN

Muestrasde frutas tropicalessalvajes y cultivadasse examinaron para determinar infestaci6n por Anastrepha spp. y Ceratitis capitata (Wiedemann)(Diptera: Tephritidae). De 440 muestras (por lo menos con 4 frutas cada una) en 201 localidadesa traves de Costa Rica, CentroAmerica, el 95%de los insectos (n = 4126)pertenecianal genero Anastrepha(A. obliqua),A. striata, A. serpentina,A. manihoti, A. pickeli, A. distincta, A. chislayae, y A. fraterculus), mientrasque 4.7%fueronC. capitata.Anastrepha spp. tuvieron una marcaadapreferenciapor plantas hospederaspertenecientes a la misma familia.A. obliquainfest6 el 92%de la infestaci6ndel mango, Mangifera indica, el 87.5%del jobo, Spondias mombin, y el 100%del yupl6n, S. dulcis, y del jocote, S. purpurea,todos estos de la familiaAnacardiaceae. A. striata se encontr6en

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Jiron & Hedstrom:Fruit Flies in Costa Rica

63

el 97.8% de las guayaba, Pisidium guajava infestadas, el 97% de P. friederichsthalianumy el 100%del gusaro, P. savanarum; estas tres especies hospederas pertenecen a la familiaMyrtaceae.A. serpentina fue la especie dominanteen plantas hospederas de la familia Sapotaceae, y se encontr6 en el 100%de todos los zapotes, Manilkaraachras infestados,el camaimito amarillo,Pouteria cainito, y el 98% del caimitomorado,Chrysophyllumcainito. A. manihotiy A. puckeliestabanasociados con casave, Manihot esculenta (45.8%y 54.2% de infestaci6nrespectivamente).A. chiclayae infest6 el 100%de las muestras de granada,Passiflora quadranqularis.A. distincta infest6 exclusivamenteespecies de Inga (Fabaceae).C. capitata fue la uinica especie de mosca de frutas encontradaen el melocot6n,Prunxus persica. El grado de infestaci6nde C. capitatafue bajo (<7%). Se presentaunafenologiaanualde las plantas hospederasmencionadas,con observacionesadicionalessobre la interacci6nentre las moscas de frutas y sus plantas hospederas.

Of about 193 species of Anastrepha found in tropicalAmerica, only a few species are of economicimportance(Norrbom1985). These are the Caribbean fruit fly A. suspensa (Loew), the South Americanfruit fly A. fraterculus (Wiedemann), the Mexican fruit fly A. ludens (Loew), the West Indies fruit fly A. obliqua(Macquart), the guava fruit fly A. striata Schiner, and the serpentine fruit fly A. serpentina (Wiedemann). Otherfruit flies infest wild plants of the tropicalwet forest. The geographicdistributionof host species of the genus Anastrephain Costa Rica has been studied twice. Salas (1958), in his publication on the Mediterranean fruit fly, Ceratitis capitata (Wiedemann), reported distributiondata from 89 locationsin Costa Rica (except for provincesof Lim6nand Guanacaste).In 1979Jir6n and Zeled6npublished anotherreport with distributional data from32 localitiesthroughoutthe country. On this occasionnine species of fruits commonly consumedby Costa Ricanswere sampled. Both studies mentionedthe possibilitythat C. capitata and Anastrephacompete for host plants. It has been shown that except on Coffea arabica L., Prunus persica (L.) Batsch,Byrsonima crossifolia L., TerminaliacatappaL., CitrusreticulataBlanco and C. sinensis (L.) Osbeck,Anastrephaappearsmore often than C. capitata in Costa Rica (Salas 1958,Foote 1967).The most commonspecies of Anastrephain commercially importantfruits are:A. striata, A. obliqua,andA. serpentina. Due to lack of quantitative information,a large scale research programwas undertakenthat focused on seasonal occurrence,geographicaldistributionand infestation rates. The present paper reports fruit fly species associatedwith host plants.
MATERIALSAND METHODS

Between April 1985 and November 1986, fruit were collected from 201 localities (Fig. 1). Each collectionconsisted of at least four infested fruits which were taken to the laboratoryand placedin wide-mouth jars as describedby Jir6n and Zeled6n(1979). Sawdustwas utilizedinstead of sand. Host plantsincludedin this study were: Psidium guajava L. (guava), P. savannarum Donn. Smith. (gudizaro), P. friedrichsthalianum (Berg) Ndzu (Costa Rican sour guava), Mangiferaindica L. (mango),Spondias purpurea L. (Spanishplum),S. dulcis Parkinson(juneplum),S. monbin L. (jobo),Manilkara achras (Mill.) Fosberg (zapote colorado), Chrysophyllum cainito L. (purple caimito),Pouteria cainito Radlk.(yellowcaimito),Calocarpummammosum(L.) Pierre (caimito),Diospyros digyna L. (tropicallegume), Manihot esculenta Crantz(cassava), and Passiflora quadrangularis L. (granada). In addition, Inga edulis Mart. I. marginata Willd. (Spanishguava), Inga Millersp., and Prunus persica (cuajiniquil),

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64
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Florida Entomologist 71(1)

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Fig. 1. Localitiesin Costa Rica where 16 species of fruits infested with Anastrepha and Ceratitis fruit flies (Diptera:Tephritidae) were collectedduring1985and 1986. (L.) Batsch (cling peach), which are not associated with commercial monoculturesin Costa Rica, were also included.Adults that emerged were pinnedfor later identification, using Steyskal'staxonomickey (1977). In cases of doubtfuldetermination,specimens were sent to Dr. A. L. Norrbom (Systematic Entomology Laboratory,ARS, USDA, Washington,D.C.).
RESULTS AND DISCUSSION

Totalnumberof adultsobtainedin the laboratory from440 fruit collectionswas 4126. Of this total 3932 (95.3%) belongedto the genus Anastrepha[A. obliqua,A. striata, A. serpentina,the cassavafruit flies A. manihoti LimaandA. pickeli Lima,the Inga fruit and 194 fly A. distincta Greene,A. chiclayaeGreene,andA. fraterculus (Wiedemann)] were C. capitata. Only0.9%of the fruitcollections were infestedby C. capitata. (4.7%) The proportionof the different fruit fly species associated with 18 host plants is shown in Table 1. Notice that infestationby C. capitata is extremely low, and except for the introduced P. persica, the figures are below 7%.The three commercially important host plants, P. guajava (n= 138), M. indica (n= 91), and P. friedrichsthalianum (n = 33), were sampledmost extensively. We foundthat P. guajava was infested by A. striata in 97.8%of all cases. Similarly,P. friedrichsthalianum,anotherspecies of Myr-

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Jiron & Hedstrom:Fruit Flies in Costa Rica

65

taceae, is also attacked primarilyby A. striata (97%).In the case of M. indica, A. obliquaoccurredmost often (92.7%).No data has been publishedon the infestationrate for wild or semi-wildAnastrepha host plants, but preliminarystudies on infestation rates for A. striata in different localitiesin Costa Rica show that as muchas 92.5%of commonguava and 97.5%of sour guava are attacked (Jir6n,unpubl.data). The flies showed a markedpreferencefor certain plant families (Table 1). An extreme example is A. distincta, which infested species of Inga (Fabaceae)exclusively and was the only fruit fly species foundattackingthe three species of Inga. Similarily, A. serpentinaappearsto prefer the family Sapotaceae.Data from Table2 supportthe hypothesisthat fruit flies prefer certainplant families.
FRUIT FLIES

Table 2 shows the annualocurrenceof adults and larvae of eight species of Anastrepha and Ceratitis associated with different host plants in Costa Rica. After a 19monthsamplingperiod, we noticedthat the phenologyof plants may vary accordingto local climatic conditions, which affect the physiology of different mango varieties (Hedstrom, et al. 1986). Many host plants are widely distributedthroughoutCosta Rica, i.e. M. indica, P. guajava and P. friedrichsthalianum,which allowedus to observe the effects of climaticpatterns on host phenology(Table2). A. obliquaWest Indies fruit fly. This species infested M. indica, P. guajava, S. purpurea,S. mombinandS. dulcis. associatedwith M. indica andSpondiasspp. (bothwild and cultivated) It was primarily were as shownin Table 1. [Of91 samplesof attackedM. indica from69 localities,92.7% infested by A. obliqua, 6.3% by C. capitata, and 0.1%by A. serpentina (Table 1)]. Individual fruits were infested by only on fruit fly species. A. obliquainfests M. incica such as from May to September,but subsequentlyinfests other hosts (Anacardiaceae) S. purpurea.S. mombinand S. purpureawhich are used by growers as living fences. with Fruits of S. purpureaare availableto female flies fromAprilto June (overlapping M. incica, Table 1). Apparently,the same applies to S. dulcis. The populationdynamicsof A. obliqua depends on the variety of M. indica (Joel et al., in press). Althoughmango varieties et al. 1985, Soto-Manitiu 1980, Guillo-Sosa was never found were not distinguishedin this study, the one knownlocallyas "criollo" infested with Anastrepha. et. Infestationof M. indica by fruit flies reaches 70%in Costa Rica (Soto-Manitiu pers. al. 1986).There is a peak shortly after the onset of the rainy season (W. Umania, comm.), which suggests that Anastrephaadults (mainlyA. obliqua)survives between cropsas pupaeunderground.In our laboratory,five adultsof A. obliquaemergedafter some 200 days as pupae in dry dirt in a jar. In agreement with previous authors, we foundthat A. obliquais the most commonfruit fly in mango(90%of infested fruits in Costa Rica, Soto-Manitiu 1986,and present in ten out of eleven varieties in Guatemala, Guillo-Sosaet al. 1984). In CentralAmericathis fly infests: Anacardium occidentale Annona spp. (an6nas),Averrhoa carambola L. (carambola), y Citrus L. (marani6n), aurantium L. (naranjaagria), C. grandis (L.) Osbeck (toronja),Dovyalis hebecarpa Warb. (kitembila), Eriobotrya japonica (Thumb.)Lindl. (nispero),Eugenia uniflora L. (pitanga),Manilkara achras (zapotecolorado),P. guajava, Spondias spp. y Syzygium 1986).We have sampledthese fruitsandfound rosa) (Soto-Manitiu, jambos L. (manzana larvae, which suggests that A. obliquaprefers M. indica, S. purpureaand S. mombin in Costa Rica.

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66
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Jiron & Hedstrom:Fruit Flies in Costa Rica

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In addition,we observedmangocultivatedfrom5 m s.m. (Jicaral,Provinceof Puntarenas) to 1300 m s.m. (San Rafael de Montes de Oca, Province of San Jose). These localities are situated in the tropical dry and rain forests, respectively. In the first region, as well as in other areas with well definedseasons (Aranjuez,Canias, Barranca, Esparza, Orotina, etc), mango is easily cultivated for commercialpurposes. On the other hand, mangois adverselyaffectedby the climatein the tropicalrain forest areas, whichhave no predictabledry season. In these areas the floweringseason of M. indica is not well defined, the pollen grains become moist and sticky after a short period of rain (Jir6nand Hedstrom1985),andthe fruits produced are heavilyaffectedby anthracnose (a fungus disease) and/orby Anastrephaspp. and C. capitata. For these reasons most orchards of M. indica are located in central Costa Rica and the Province of for commercial Guanacaste,whichrepresentthe most suitableareas climatically mango production(Fig. 2a). Varieties of M. indica differ in their fruitingphenology,even in the same locality(Soto-Manitiu et al. 1986).Consequently M. indica fruits are available from Marchto September,in many differentlocationsof Costa Rica. A. striata guava fruit fly. The range of the guava fruit fly is the widest in the Anastrepha group, from in the north (Hedstrom1985)to Ecuadorin the south (Hedstrom1987).This Guatemala to the naturalrange of Psidium (Myrtaceae), corresponds whichthe fly caninfest under many climaticconditions(voltinismvaries widely as does the phenologyof P. guajava). We foundthis species from sea level (PlayaJunquillal,Provinceof Guanacaste) to 1510 m s.m. (Monteverde,Provinceof Puntarenas). In 138 samples of P. guajava from 94 localities 97.8%were infested by A. striata, P. guajava, is infested occasionallyby A.obliqua (0.7%),A. fraterculus (Wiedemann) (0.7%)and C. capitata (0.7%)(Table 1). A. striata infests P. guajava year-round,however, populationdynamicsdependon the geographicarea. Recently we found that in Guapiles,Province of Lim6n(tropical wet forest), A. striata survives year-roundin secondaryhosts, i.e. Persea americana L. (avocado)(Lauraceae),P. friedrichsthalianumand S. dulcis. As mentionedbefore, our samples showed that A. striata infests primarilythree species of Myrtaceae.A. striata was also foundin two other secondaryhosts, S. mombin and C. cainito (Table 1 and 2). Commonguava may be the most widely distributedAnastrepha and/or Ceratitis host plant in Costa Rica. It is most abundantin tropicalrain forest on Caribbean facing slopes. These do not have a pronounced dry season (Fig. 2b). Due to genetic variability, fruits supportpopulationof A. striata throughoutthe year. Table 1 shows that about 98%of the fruit flies on commonguava are A. striata. Occasionally we foundisolated guava trees resistant to infestationby A. striata. On the other hand, sour guava or "cas"has a restricted range in Costa Rica. It is not grown commercially due to high fruit fly infestation. Our data come from isolated but commercially exploitedtrees, since we knew of no large orchards.Consumersusually acccept these fruits even when a few larvae (that can be removed manually)are present. A. serpentinafruit fly. We collected73 samples,fromM. achras, C. cainito, C. nammosum andP. cainito, with infestationrates of 98-100%. ApparentlyA. serpentina prefers Sapotaceaefruits in the tropicalwet (Provinceof Lim6n)and dry (Provinceof Guanacaste) forests. The Sapotaceaefruits were infested primarilywith A. serpentinain wet and dry zones. In

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70
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March, 1988

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Jiron & Hedstrom:Fruit Flies in Costa Rica

71

Guapiles(Provinceof Lim6n),adultflies appearwith fruits of this plantfamily,showing a close correlationwith the phenologiesof C. cainito and P. cainito. A. serpentina was captured in traps set in a commercialorchardof M. indica in Orotina,Province of Alajuela, duringthe fruitingof Sapotaceae,at the end of the dry season and duringthe rainy season (Table2). We caught 24 adults, but none emerged from fruits culturedin the laboratory,which only producedA. obliqua. This suggests that A. serpentina feeds on mango without ovipositingin the fruit or that the species oviposits but does not survive. Species of low incidence. A. fraterculus (on P. guajava) is an economicallyimportant fruit fly in South America(Malavasiet al. 1980).We foundadults only twice, in P. guajava (Santiagode San Ram6n,Provinceof Alajuela, and La Selva BiologicalStation, Sarapiqu,Province of Heredia). The cassava fruit flies, A. pickeli and A. manihoti, apparentlycoexist on M. esculenta through spatial separation:the former in terminal buds, the latter in fruit capsules(Saunders& Salazar1979).We have capturedthe passiflorafruit fly A. limae, in McPhailtraps in areas where M. esculentais not cultivated(Provinceof Guanacaste), which suggests that they also develop in wild euphorbs. The latter species was also collected for the first time in Costa Rica while foraging on P. quadrangularisand reported (together with A. pickeli) (Hedstrom et al. 1985). Similarly, A. chiclayae appearedin the tropicalwet forest in La Selva BiologicalStation(Provinceof Heredia), and in Buenos Aires de Puntarenas(Provinceof Puntarenas)where the host P. quadrangularis was not found (it occurs in La Garita, Province of Alajuela, where the associationswas found)(Table 1). The Inga fruit fly A. distinctais closelyassociatedwith Fabaceae(Table1), including wild species and those used to provide shade in coffee plantations. The female can ovipositin a woody fruit, seldominfested by other tephritids. Its range seems to match the distributionof the host. We captured C. capitata in an orchardof M. indica during 1984 (Fabio Baudrit Experimental Station, La Garita, Province of Alajuela) and found large numbers of female of the species in the traps early in the mangocrop season (Hedstromand Jir6n 1985).The presenceof C. capitata coincidedwith the fruitingof tangerinsin La Garita. Apparently, in Costa Rica, C. capitata is the only fruit fly infesting tangerines and oranges (Fischel 1982). Although we have collected over 11,000 adult Anastrepha in Costa Rica duringthree years of sampling(Jir6n& Hedstrom,unpubl.data), we have not foundA. ludens. If it occuredhere, it wouldprobablyhave displacedCeratitisfrom citrus species (Salas 1958, Foote 1980),althoughthis hypothesisneeds to be tested. C. capitata exists at low levels in Costa Rica and does not cause enough economic damageto justify control(Table 1). For example, on coffee C. capitata infests only 5% of the berries and causes no importanteconomicimpact (Fischel 1982). Apparently, coffee sustains the population levels. The importhroughoutthe year at low population tance of C. capitataas a fruit pest in Costa Ricais related to the wide range of potential host plants, which includethose for export and hence present quarantinedifficulties.

Fig. 2. Distributionof fruits. (A) Mango(Mangiferaindica L.) collections,mainly infested by the West Indies fruit fly Anastrepha obliqua (Macquart), primarilyin the provinces of Guanacaste, Puntarenas and Alajuela, in the transitional tropical dry forest. (B) Commonguava (Psidium guajava L.) collections, mainly infested by the in the tropicalrainforest life zone. guavafruitfly Anastrephastriata Schiner,primarily

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72

Florida Entomologist 71(1)

CONCLUSIONS

March, 1988

Most species of Anastrepha have ranges that match their hosts. For fruits widely distributed in Costa Rica, the most common fruit flies and host plant families are: A. obliqua on Anacardiaceae, A. striata on Myrtaceae, A. serpentina on Sapotaceae, and A. manihoti and A. pickeli on Euphorbiaceae. Of low incidence are C. capitata (on P.

guajava, P. fiedrichsthalianum,M. indica, M. achras, and C. cainito). Four samples

of P. persicae were completely infested by C. capitata. The apparent correlation between species of Anastrepha and host families suggests a biochemical relationship, and the voltinism of part of these populations seems to depend on the phenology of the primary host plant. Probably, many unknown wild hosts of the appropriate families play an important role in the population dynamics of the flies.
ACKNOWLEDGMENTS

This research was supported by the Consejo Nacional de Investigaciones Cientificas y Tecnol6gicas (CONICIT) de Costa Rica and the Vicerrectoria de Investigaci6n de la Universidad de Costa Rica. We are grateful to R. Zeled6n, Minister of Science and Technology, Costa Rica, J. M. Soto-Manitiu, J. Monge-Najera, M. J. West, R. G. Mexz6n and H. P. Sauter at the Universidad de Costa Rica, A. L. Norrbom, Systematic Entomology Laboratory, USDA, ARS, Washington D.C., and three anonymous reviewers for comments on the manuscript. The authors thank H. Lezama, I. M. Gonzalez, L. E. Cordero and M. Diaz, Universidad de Costa Rica, for assisting in the field work. REFERENCES CITED FOOTE,R. H. 1980. Fruit fly generasouthof the United States (Diptera,Tephritidae).

USDA Tech. Bull. No. 1600: 1-79. en la FISCHEL, M. 1982. Fluctuaci6nen la densidadde poblaci6ny parasitoidismo mosca del Mediterraneo (Ceratitiscapitata) (Diptera, Tephritidae)en frutos de
cafe (Coffea arabica) en la regi6n de Santo Domingo de Heredia, Costa Rica.

Tesis, Fac. Agron., Universidadde Costa Rica, 53 p.

GUILLO-SOSA,M., F. M. ESKAFI, AND F. J. MANUEL. 1984. Identificaci6n de es-

pecies del genero Anastrepha, sus enemigos naturales y su preferenciaa diferentes variedadesde mango en el Departamentode Retalhuleu, Guatemala. Tikalia3(1): 15-27. HEDSTROM,I. 1985. Anastrepha striata (Diptera, Tephritidae),new to Guatemala. Rev. Biol. Trop. 33: 195-196.
. 1987. Fruit flies (Diptera, Tephritidae) infesting common guava (Psidium

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TOXICITYAND LETHAL TIME OF N-ETHYL PERFLUOROOCTANESULFONAMIDE AGAINST TWO SUBTERRANEAN TERMITE SPECIES (ISOPTERA:RHINOTERMITIDAE)
NAN-YAO Su AND RUDOLF H. SCHEFFRAHN Ft. LauderdaleResearch and EducationCenter University of Florida, IFAS 3205 College Ave. Ft. Lauderdale,FL 33314

ABSTRACT The topical LD50of N-ethyl perfluorooctane sulfonamide (GX071)was estimated at 9.94 Rg/g against the Formosansubterraneantermite, Coptotermes formosanus, and 68.61 Rig/gagainst the eastern subterraneantermite, Reticulitemnes flavipes. Under force-fed conditions, C. formosanus was ca. three fold more susceptible to GX071 = 4.22 ppm)than R. flavipes (LC50 = 13.6 ppm).Whenappliedtopically,5-15days (LC50 elapsed before 90%of the R. flavipes died (corresponding dose range:100-250 ,ug/g), while a similarmortalityrate was recordedfor C. formosanus after 2-7 days exposure at a lower dose range:14.0-37.5,ug/g.Ninety percentof bothtermite species were killed 3-12 days after being force-fedwith GX071,but at a lower concentration range for C. formosanus.
RESUMEN

El LD50tipico de N-ethyl perfluorooctane sulfonamidae [G X 071) se estim6 ser 9.94 ,ig/g contra la termita subterraneade Formosa, Coptotermes formosanus, 68.61 ,ug/g

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