You are on page 1of 24



Quick links to online content
Annu. Rev. Entomol. Copyright

© 1994

1994. 39:155-78

by Annual Reviews Inc. All rights reserved

Annu. Rev. Entomol. 1994.39:155-178. Downloaded from by University of Guelph on 10/10/12. For personal use only.

M. Aluja
Instituto de Ecologfa, A.C., Apartado Postal 63, 91000, Xalapa, Veracruz, Mexico
Diptera, Tephritidae, biosystematics, zoogeography, biology, ecology, life strategies, behavior, pest management

Flies of the genus

Anastrepha (Diptera: Tephritidae) are among the world's

most devastating agricultural pests. At the same time, they display remark­

able ecological and behavioral characteristics, which have served as models

in the development of general theories on insect mating systems and the physiology of host marking.

Anastrepha species are endemic to the New

World and restricted to tropical and sUbtropical environments. The genus's most of the Caribbean Islands. There are range extends from the southern US to northern Argentina and includes

The great majority of these are poorly known biologically, and knowledge is basically restricted to seven economically important species:

184 described Anastrepha species.

Jraterculus, grandis, ludens, obliqua, serpentina, striata, and suspensa. Much of what we know today about Anastrepha is based on very thorough studies of its basic biology carried out between 1900 and 1944 (13, 53, 85, 115, 188; see also 21, 45, 167). More recently, work has concentrated on phylogeny
and taxonomy, behavioral ecology, chemical ecology, demography, and species surveys. and technically stagnant for the past

McPhail traps are still standard procedure even though such methods have been proven ineffective and environmentally questionable. literature on

Anastrepha control has in many ways been conceptually 35 years: poisoned bait sprays and

Here, rather than providing an all-encompassing review of the voluminous

Anastrepha, I attempt to expose the reader to a balanced

155 0066-4170/94/0101-0155$05.00



selection of poorly advertised studies from Latin America as well as work published in mainstream journals. I place special emphasis on recent ad­ vances in phylogeny, taxonomy, distribution, host-plant relationships, de-' mography, population genetics, behavioral ecology, and some new control methods and management approaches.

Annu. Rev. Entomol. 1994.39:155-178. Downloaded from by University of Guelph on 10/10/12. For personal use only.

Taxonomy and Phylogeny
Anastrepha belongs to the family Tephritidae (true fruit flies) and was recently placed in the tribe Toxotrypanini of the subfamily Trypetinae (67). Many species of Anastrepha were first described in other genera such as Dacus, Pseudodacus, Lucumaphila, Phobema, and Trypeta (132,185). To date, the most thorough revision of the genus is Stone's 1942 monograph (188), which was partially updated by Steyskal (184), Zucchi (196), Foote (66), Norrbom (132), and Hernandez-Ortiz (83). One should note that the taxonomy of Anastrepha is based almost exclusively on the adult female. Only recently have efforts been directed toward identifying males (132). The immature stages of Anastrepha are poorly known. So far, descriptions of third-stage larvae have appeared for only the following species: A. bistrigata, A. consobrina, A. distincta, A. Jraterculus, A. grandis, A. interrupta, A. limae, A. ludens, A. nunezae, A. obliqua, A. pal/ens, A. sagittata, A. serpentina, A. striata, and A. suspensa (13, 23, 42, 49, 64, 75, 82,132, 142, 181-184, 194). Eggs have been described for the following species: A. atrox, A. cordata, A. leptozona, A. iudens, A. nigrijascia, A. obliqua, A. pal/ens, A. pittieri, A. serpentina, A. striata, and A. suspensa (57, 98, 132).
Cladistic analysis and immunological data support the placement of

Anastrepha in a monophyletic group with the genus Toxotrypana (67, 93, 132). Anastrepha is most probably of South American origin (83). Morgante et al (119) suggest that many species of Anastrepha are relatively young
and that speciation has been rapid and recent. The first serious attempts at studying the phylogenetic relationships of Anastrepha were recently pub­ lished by Norrbom (132-134) and Norrbom & Kim (136). These authors divided the genus into 18 preliminary groups mainly based on morphological

benjamini, chiclayae, cryptostrepha, dacijormis, dentata, doryphoros, !raterculus, grandis, leptozona, mucronota, pseudoparaUela, punctata, schausi, serpentina, spatulata, striata, ramosa, and robusta. Given
characters: the size of the group, the rate at which new species are being described and the presence of many cryptic species, future infrageneric classifications will have to be based, in my view, on a combination of criteria such as

although A. cordata. Its range covers part of North America (southern Florida. and further knowledge of the ecology and behavior of the group. A. and at least some species in the A. Certain species (e. 4% are found only in North America (Mexico and the US). crebra. eggs are laid singly or in clutches. Downloaded from www. and Argentina (25) reveal that the range of species such as A.ANASTREPHA B IONOMICS AND MANAGEMENT 157 morphology.or mesocarp region of ripening host fruit. Nevertheless. The larvae of most species feed exclusively on the fruit pulp. bahiensis. A thorough zoogeographic analysis at this stage is limited by the fact that the exact distribution of most species of this genus has not yet been established. and A. 15% are restricted to Central America (including Panama). cytogenetics. A. sagittata). and A. 126. serpentina. A. and most of the subgroups are widespread. the Rio Grande Valley in Texas. Larvae go through three instars before leaving the fruit and burrowing into the ground to pupate (on occasion larvae pupate inside the host fruit). Peru (95. Central and South America (except Chile and southern Argentina). Costa Rica (92). A. A. For example. For personal use only. obliqua. obliqua lays one egg per oviposition (43) whereas A. cryptostrepha groups feed on pulp and seeds or only on . A. Rev. A. acris. 83). 43% are only found in South America. Some species have an extremely slender and long ovipositor (e. 1994.g. BASIC BIOLOGY Life Cycle and Life Strategies The basic life cycle is very similar among all Anastrepha species for which the biology is known. Venezuela (41). grandis can lay clutches of up to 110 eggs (52). Brazil (40. Overall. and almost all of Mexico).org by University of Guelph on 10/10/12. Depending on the species. A. 96). A. according to Norrbom & Foote (135) the distribution at the species group level in Anastrepha lacks a discernible pattern. and 1 % are found only in the Lesser and Greater Antilles (84). 196. Annu. bicolor. which may allow them to oviposit into seeds instead of the pulp (13.annualreviews. 132). A.g. of the 184 Anastrepha species described so far. striata is remarkably large. In the majority of species. A. Zoogeography Anastrepha is endemic to the New World and is restricted to tropical and subtropical environments (83).39:155-178. and most of the Caribbean Islands (84). jraterculus. tumida) are associated with tropical rain forests (83). Regional studies in Mexico (13. leptozona. A. 197). isozyme studies. molecular (DNA) analyses. tripunctata) seem to be associated with tropical deciduous forests with long dry seasons. A.g. while others (e. manihoti feed on stems and buds (140). the females deposit their eggs in the epi. alveata. daciformis. Entomol.

165) and rearing medium (43. and is sexually dimorphic (173). Host-plant records are only known for approximately 39% of all reported . Anastrepha species typically inhabit highly variable environments(i. Entomol. crebra. 99. strictly monophagous species such as A. 1 15).annualreviews.158 ALUJA Annu. and patchy or isolated in space) where they live in close association with their host plants. adults crawl to a sheltered spot nearby until their wings unfold and dry. Hernandez. Host-Plant Relationships Norrbom (132) indicates that Anastrepha is clearly not old enough to have strictly coevolved with its hosts but suggests that host-plant associations of Anastrepha appear to be at least partially correlated with phylogenetic relationships within the genus. zeteki. Exit of larvae from fruit is determined by fruit characteristics (e. Zw6lfer (198) has indicated that they are opportunistic. 114. dentata. An interesting feature of Anastrepha host-plant relationships is the close association between certain species and particular plant taxa: for example. 115).39:155-178. pH. multivoltine. For personal use only. 53. 1994. personal communication). Rev. Before becoming sexually active. Pupation depth is greatly influenced by soil pH and type (e. For example. internal temperature. by University of Guelph on 10/10/12. unpredictable. polyphagous species. robusta. the chiclayae and pseudoparallela subgroups are associated almost exclusively with the genus Passijlora (Passifloraceae). but we simply know too little about Anastrepha field ecology to utilize hard-and-fast categories at this stage. ludens or A. clay) and the degree of soil compaction and humidity (28. obliqua.e. and are usually long-lived. 100). Downloaded from www. and leptozona subgroups are mainly associated with the family Sapotaceae (132). may be able to cope with resource patchiness and ephemerality by going through periods of estivation and by tracking seasonal hosts scattered over large areas by moving effectively from habitat to habitat(5). sand.g. After emergence. in which adult emergence is closely timed with host-plant fruiting phenology. and highly mobile. adults go through a maturation period during which they need to feed regularly on carbohydrates and water to survive and on protein sources to allow for gonad maturation (45). In contrast. 137. daciformis. seasonal. exhibit high reproductive potential. are probably univoItine and undergo obligatory diapause(V. or ephemeral in time. The rate of immature-stage development is determined to a great degree by environmental factors such as temperature (14. These attributes are broadly applicable to the genus as a whole.g. The fact that the cryptostrepha and daciformis groups (primitive Anastrepha) are strongly associated with sapotaceous hosts suggests that the Sapotaceae may have been ancestral hosts for Anastrepha (132). degree of rotting) and by physical signals experienced when ripe fruit drops to the ground or is exposed to rain (53. seeds (13. 188). The serpentina. such as A.

annualreviews. For personal use only.39:155-178. Rev. 76. carbohydrates. 13). Unfortunately. minerals.ANASTREPHA BIONOMICS AND MANAGEMENT Anastrepha species 159 (137). and water for development. leaf and fruit surfaces. Furthermore. bird feces. 116. but optimal male salivary gland development (important for sexual pheromone production) and female ovary development depend on extrinsic sources of protein (62. As is the case with other tephritid genera. 137). stenophagous. or are related to cultivated and. 68. seven economically important spe­ cies account for -70% of all reported Anastrepha host records (132). vitamins. When the larvae feed on seeds. Laboratory studies with A. Such nutrients are obtained in nature from liquids oozing from overripe or damaged fruit. Adult flies can survive for long periods of time on carbohydrates alone. many host-plant associations are a local phenomenon (i. 162). is confounded by the fact that Annu. cordata. For example. Nutrition As is the case with other tephritids. and survival (76). a mixture of all amino acids at a concentration of 15% mixed with vitamins and sucrose yields the greatest number of eggs and the highest egg eelosion percentages (27).org by University of Guelph on 10/10/12. exotic hosts (8. as in A. reproduction. in many cases. G Siller & R Rosiles. Almost no information is available on wild host plants in unperturbed environments. The metabolic pathways used by larvae to detoxify many deleterious compounds found in food plants are poorly understood. Anastrepha have anatomical adaptations . immature and adult Anastrepha require amino acids. which contains alkaloids close to the virulent toxin strignin (M Aluja. 113) have found alcohol dehydrogenase (ADH)-specific activity in larvae of several Anastrepha species and suggest that selective pressures for enhanced ethanol metabolism have occurred in the evolutionary history of Anastrepha species. Matioli et al (112. unpublished data). 1994. Such clas­ sification. oligophagous. Entomol. acris feed on Hippomane mancinella fruit (Euphorbiaceae). a considerable number of these records are incidental or the result of misidentifications. were obtained under laboratory conditions. Furthermore.e. Based on host-plant lists. Anastrepha species have been classified as mo­ nophagous. female and male gonad development is completed shortly after emergence and apparently without the need for protein intake (M Aluja & V Hernandez. and polyphagous (2). Downloaded from www. Associations with Bacteria The· role of bacteria in the nutrition and survival of tephritid larvae and adults is best known in Bactrocera and Rhagoletis species (65). though useful in general terms. larvae of A. and rain drops (5. ob/iqua have shown that the optimal protein/carbohydrate ratio for ovarian development is 3:5 (63). polyphagous species behave as monophagous or stenophagous species in certain areas). unpublished data).

ludens and A. obliqua. fraterculus. Population genetics studies in Anastrepha have been restricted to one species: A. Entomol. and exhibits greater morphological variation than related species. and Pseudomonas ovalis were also identified. jraterculus = .160 ALUlA Annu. 1994. striata (36. suspensa. Steck (180) carried out an isozyme analysis that revealed sharp genetic discontinuities among A. bistrigata.. Proteus ratgeri. by University of Guelph on 10/10/12. the bacteria Micrococcus sp. ludens and A. Bacteria housed in the gut of Anastrepha and other fruit fly larvae and adults are known as fruit fly type bacteria (Enterobacteriaceae) (65). Karyotype analyses have been performed on A. Recently. Pseudomonas sp. Fletcher. Pseudomonas jluorescens. 118. Proteus mirabilis. it represents an unresolved complex of cryptic species (13. bistrigata. Klebsiella ozanae. A. widely distributed. A. In a previous study on A. These studies have revealed that the typical chromosome number in Anastrepha is 2n 12. 176). This species is polyphagous. According to several authors. A. but such findings could have been the result of artificial laboratory conditions (26. pseudoparallela. A. Enterobacter gergoviae. What role bacteria play in the biology of adult flies is still poorly understood. 156).39:155-178. pickeli. fraterculus. pickeli. 118). 180.defense chemicals.. Anastrepha adults regurgitate food on leaf surfaces shortly after feeding on fruit juices (6. and Pseudomonas cepaceae from the gut and eggs of A. Erwinia sp. A. in the break· down and digestion of fruit tissues. and in suppressing pathogenic fruiHot microorganisms (65). Enterobacter agglomerans. jraterculus. and A. Rev.annualreviews. Four distinguishable karyotypes in Brazilian popula· tions have been described (176). serpentina. for housing microorganisms.X2 Y mecha· nism of sex determination (177). but no host·dependent genetic differentiation is evident( lOS. Variations in the diploid number have been detected in a few species.. A. in detoxifying plant. Murillo et al (123) and Solferini (175) have isolated bacteria such as Acinetobacter lwojfi. obliqua. Downloaded from www. particularly those having an XX:XO or X1X. Bacterial syinbiotes may also play an important role in synthesizing essential amino acids. Proteus vul· garis. A. such as gastric ceca in larvae and an esophageal diverticulum (bulb) in adults (175). Klebsiella pneumoniae. 106. suggested that flies inoculate the surface of fruit and utilize the resulting bacterial colonies as protein sources (65 and references therein). Klebsiella oxytoca. A.X:zX2:X. interpreting a similar behavior in Bactrocera spp. POPULATION ECOLOGY Population Genetics Both the cytogenetics of individual species and population genetics of Anastrepha have been studied. 119. A. 10). Enterobacter cloacae. For personal use only. 188).

gross and net fecundity) have been determined for A.: Bracana­ strepha anastrephae. A. O. Scattered efforts by early workers are incomplete and were carried out under laboratory conditions (13. Longevity and survivorship curves have also been described for A. Costa Rica (91. craw­ Jordi. and Pachycrepoideus vindemiae (Pteromalidae). (Diaprididae). Surveys in Mexico (9). 124) indicate that the following indigenous hymenopterous parasitoids commonly attack Anastrepha spp. in large vs small (mean pupal weight of 24 vs 18 mg.g. serpentina. preadult survival and development rates. coastal Venezuela. suspensa under laboratory conditions (43). ' Basic demographic parameters (e. Rev. but the true impact on the population dynamics at the field level has never been determined. brasiliensis have . 53. Opius hirtus. Steck (180) concludes that there is strong evidence that a complex of cryptic species is included in the nominal species A. 193). O. larvae. For example. Costa Rica. D. Food type. Eucoila pelleranoi (all Cynipidae). and fly density influence life expectation and gross fecundity rates (101. ludens individuals can live as long as 11 and 16 months respectively. All immature stages (eggs. populations from northeastern Brazil. Aceratoneuromyia indica (Eulophidae). brasiliensis. Odontosema anastrephae. mean gross fecundity was 1597 vs 1450. 1994. adult survival. and A. bistrigata (29). For example. ludens. including Diachasmimorpha longicaudata and Biosteres arisanus (Braconidae). vierecki.g. 101). O. D. Entomol. Two points stand out from these studies: (a) gross fecundity rates and daily egg production under ideal laboratory conditions can be extremely high (e. availability of water. Few studies in the vast Anastrepha literature examine the causes of mortality. and Mexico were all very similar. obliqua. Downloaded from www.annualreviews. populations from southern Brazil. pupae) of Anastrepha are attacked by a series of indigenous parasitoids. Andean Venezuela. Jraterculus. sororcula and A. aerolatus and D. Age at first reproduction is strongly influenced by temperature and ranges from 8 to 20 days depending on the species (13. For example. argentinus (all Braconidae). Note that D. Demography and Population Dynamics Annu. (Eucoilidae). 115). zeteki. and Psylus sp. Guatemala (60). and Argentina (61. 43. A. respectively (101). Adult survivorship curves of several species studied so far are close to type III curves. respectively) by University of Guelph on 10/10/12. 53) and 12 months under field conditions (169). 1000 eggs/female) and (b) adults can live for prolonged periods of time. ludens.39:155-178. Cothonaspis sp. under laboratory conditions (13 .ANASTREPHA BIONOMICS AND MANAGEMENT 161 populations. Doryctobracon aerolatus. and Peru were ge­ netically distinct from the first group and possibly from each other as well. For personal use only. Some exotic parasitoids are also quite commonly encountered. Ganaspis carvalhoi. Biotic mortality factors have been repeatedly identified. female and male A. 115). fly size. Brazil (12). D. tucumanus. In contrast.

ludens (143). At lower elevations (680 m). of 12 species captured. host plant species richness and diversity) and by altitudinal gradients. The most important abiotic mortality factors regulating population dy­ namics are water and temperature. Belonuchus ru fipennis) and ants (e. Belize (88). e. 178). Rev.39:155-178. Venezuela ( l09).org by University of Guelph on 10/10/12.g. ludens. 13) that attack both larvae and pupae. a picture is emerging indicating that adult populations in commercial orchards exhibit strong fluctuations from year to year and that these fluctuations are correlated with two factors: availability of host plants and climatic factors (especially rainfall). Entomol. and may reduce survivorship (13). Xenophygus analis. oviposition.162 ALUJA Annu. Colombia (138).70. Too much or too little water causes both immatures and adults to die. only 4% of individuals were A. BEHAVIORAL ECOLOGY Diel Rhythms of Activity and Resource Utilization Patterns Under laboratory conditions. Downloaded from www. one or two are dominant. 30% of all individuals were A. 1994. feeding. been recovered from wild. striata adults collected in the field (79). In monocrop orchards. Excess water also hastens emergence from the fruit and concomitant puparium formation.and second-instar larvae to die. 108). Adult Population Fluctuations Based on studies in the US (131). in mixed mango orchards at 1100 m above sea level. Other biotic mortality factors are staphylinid beetles (e. g. . suspensa larvae have been shown to feed within fruit continously over a 24-h period (189). Fruits exposed to rain decay faster and cause fIrst. Pupal desiccation in dried soil appears to be a major mortality factor. For personal use only. Solenopsis geminata) (2. fluctuations are probably dampened by the presence of alternative host plants. For example. Trapping studies in orchards clearly show that even though up to 15 Anastrepha species are commonly captured. The first days after puparium formation are critical in this respect (115). population numbers typically reach a peak shortly after host fruit has ripened and crash when no hosts are available. Costa Rica (90. In mixed orchards. of 14 species captured. Excessively dry conditions may reduce female fecundity and affect survival rate of newly emerged adults. For example in Mexico. 144). Mexico (2. mating. and Brazil (104. The degree of species dominance is influenced by ecological background (i.annualreviews. A.e. Similar results have been reported for Costa Rica (90) and Brazil (126). Pathogens have also been described. Behavioral partitioning of adult daily activities in Anastrepha (i. 126). Stigmatomyces aciurae fungi (ascomycetes) have been reported on A. native host plants in unperturbed environments (12.

flies exhibit plasticity and can adapt to local microhabitat conditions (5). striata. Shaw et al (169) trapped tepa-sterilized A. A. A. pseudoparallela (6. For personal use only. ludens. and others. Some species only call (Le. Trivial Movements and Migration LARVAE Movement of larvae is related to the maturity of the host fruit. I-day-old flies had a significantly greater mean flight propensity than older flies (15. 62. Downloaded from www. which usually takes place during morning hours. 13. ludens fly -135 km from breeding sites in Mexico to invade citrus groves in southern Texas. Other behaviors seem to follow species-specific patterns. in A. Christenson & Foote (45) have described long-range displacements and report that A. individual flies differed significantly in their mean flight times when measured repeatedly over periods of 9-10 days. oviposition was observed at precisely the period of time avoided under environmentally harsh conditions (5). in females the time between the shortest and longest flight increased 30-fold (15). 146). For example. In citrus. suspensa (5-7. if members of the same population were transferred to a more sheltered environment. Under laboratory conditions. Furthermore. obliqua call both during the morning and afternoon (5). grandis. serpentina. such as A. Also.ANASTREPHA BIONOMICS AND MANAGEMENT 163 Annu.7% emerged between 0600 and 1000 hs (115) and in A.3% emerged between 0900 and 1200 hs (10). ADULTS Research on Anastrepha adult movement includes laboratory stud­ ies of tethered A. For example. ludens individuals aimed at establishing differences between irradiated and unirradiated or between wild and labo­ ratory-reared insects (159). Entomol. robusta (4). 1994. obliqua. ludens up to 36 km from their release by University of Guelph on 10/10/12. An exception to this seems to be time of adult emergence. ludens. obliqua females were never seen ovipositing in Spondias purpurea fruit during midday when ambient temperature could reach 45°C. suspensa larvae hatch in the flavedo (the part of the fruit where eggs are deposited) and move to the albedo and pulp as they grow and the fruit ripens (39). as well as field studies aimed at establishing short (intraorchard) and long-range (interorchard) displacement patterns. 107). and resting) is quite different in the few species where information is available. However. 52. A.e. emission of a series of courtship sounds through wing fanning and release of a sexual pheromone) in the early morning. 44. 44. resting and oviposition. Jraterculus. Rev. 80. A. 160).39:155-178. and A. For example. i. A. 10. A. . In the case of behaviors such as feeding. suspensa and A. 95. such as A. and A. Other species such as A. A.annualreviews. ludens. Work on this aspect of Anastrepha biology has been marred by the lack of a theoretical framework. A. striata. restrict their calling activities to the late afternoon.

At the orchard level. 10. Rev. For personal use only. Entomol. Such patterns have been documented using release-recapture methods (58) and through direct obser­ vations (5). A. ludens. and by University of Guelph on 10/10/12. they will not leave the release site (145). Downloaded from www. 1994. obliqua.3 0). Apparently both olfactory and visual stimuli lead gravid females to suitable oviposition sites. suspensa (6. striata. and the literature is rife with anecdotal accounts about the "migratory" (3). Regurgitation behavior has been reported in A. These essential nutrients were made available by the plant through leaching and guttation processes. striata. In studying (Rhagoletis pomonella). A.39:155-178. and oviposition substrates are plentiful. water. Females typically land on fruit. bubbling (formation of a drop of liquid of varying sizes at the tip of proboscis while fly sits motionless). serpentina. release-recapture studies with A. In my opinion.5 drops and reingest them within 12 min (10). Hendrichs and collaborators (81) determined that through grazing on leaf surfaces adult flies could accrue small amounts of carbohydrates and some dabbing (or grazing) behavior in another tephritid fly proteins. the use of the term migration should be avoided unless threshold responses by flies to vegetative stimuli are tested. sucking (absorbing water droplets or liquid oozing from a fruit). A. food. Oviposition Anastrepha is a dynamic process that follows a (149). fraterculus. Anastrepha speci es such as A. and regur­ While feeding. Within a certain area. 17). lack of host plants. Factors influencing final choice are poorly understood. inspect it by walking on the surface (exhibiting Oviposition behavior in pattern closely resembling other frugivorous tephritids . or adult food sources) they will quickly leave the release site (58. A. adult gitating (deposition of a series of regurgitated drops on a leaf or fruit and reabsorption of those drops after varying intervals of time) (10). dry conditions. Overall fly mobility after release is low (16.80. but these can also be inferred from trapping studies that show that most flies are captured in border rows (2). ludens are known to move back and forth between native vegetation and orchards or between two capacity of adult flies habitats that harbor two sets of essential resources. Feeding Anastrepha flies exhibit the following behaviors: dabbing (repeated lowering of proboscis to touch the surface of leaves while walking at increased rates of turning).annualreviews. ludens show that wind affects displacements of flies (mean fly movements are oriented to directions similar to those of the prevailing wind) (16. individuals deposit a mean of 23. obliqua and A. If flies are released in a place where vegetation. obliqua or A. but it is strongly correlated with environmental conditions. 169). But if flies are released in an unfavorable environment (e. In A.164 ALUlA Annu.175).g.

the process of female choice can take up to two hours. repeated head butting bouts). and A. 107). suspensa (7. occur (10). Within a lek (or outside of it) males typically emit a series of courtship and aggressive songs via rapid wing vibrations (190. Work with A. 174). serpentina. control strategies can be summarized as follows: placement of McPhail traps. Mating Anastrepha males follow two distinct mating strategies: (a) resource defense polygyny exhibited by species such as A.annualreviews. application of poisoned bait sprays for monitoring and control. 55.ANASTREPHA BIONOMICS AND MANAGEMENT 165 Annu. ludens. 191) and release a sex pheromone through the mouth and anus (127. HMPs actually seem to stimulate continued host-marking behavior after egg laying (139). fraterculus. bistrigata that are strictly mo­ nophagous and patrol and defend clumps of fruit that are attractive to receptive females (121) and (b) lek polygyny exhibited by polyphagous species such as A. sounds. Downloaded from www. obliqua. 120. The males make a precopulatory sound as they mount the back of a potential mate or if a female becomes restless during copulation (172). Although they deter females prior to egg­ laying. in my opinion. MANAGEMENT Anastrepha control and management has. including repeated mating attempts and labellum-to-Iabellum contacts (during which actual trophalaxis takes place). 130). A. A. obliqua. females deposit a water­ soluble host-marking pheromone (HMP) after ovipositing (147. with clutch size strongly affected by host size (24). Males deposit pheromone on leaf territories and defend them with a mixture of postures. Rev. striata). preferring larger males (35). Pheromone release is achieved by puffing expanded pleural abdominal membranes and an everted proctiger (129). Larger males sing better-quality songs and presumably pass more sperm to females (172). A. bore. ludens has suggested that host-marking is regulated by sensory adaptation or habituation to HMP in conjunction with dosage-dependent restoration of inhibition of the motor pattern (139). For personal use only. elaborate close-range courtship displays and interactions. personal communication). 33. F. ludens. eggs are deposited singly or in clutches. suspensa.39:155-178. A. males establish territories preferably on the underside of leaves of host and nonhost trees (5.physical encounters ( by University of Guelph on 10/10/12. fraterculus. Diaz. 148. striata. A typical characteristic of the Anastrepha lekking system is that females make discriminating mate decisions. A. From a historic perspective. 170). and . been conceptually and technically stagnant for the past 35 years. and . 80. and oviposit. In the few species studied so far (A. A. In species such as A. As discussed earlier. In lekking species. 1994. Entomol. During this time. A.

103. obliqua. designing color-based traps. 186). has several drawbacks: it is expensive. ludens. For example.39:155-178. suspensa) when used in spherical. ludens. A. fraterculus. suspensa females (73). 73. Colors such as yellow and orange. suspensa females. Downloaded from www. Rev. For example. high densities of traps are needed to detect them (38). we (6) found that out of 665 flies that landed on the exterior of the trap. it preferentially captures females (88). Adult Detection Mechanisms Without question the most widely used traps to monitor and in some cases control Anastrepha populations are glass and plastic versions of the McPhail trap. rectangular or cylindrical traps (31. Human urine has been found to be 10 times more effective than torula yeast in attracting A. serpentina. laboratory reared A. . 1994. 111. Recent work on increasing the effectiveness of Anastrepha monitoring mechanisms has progressed along three lines: developing alternatives to standard hydrolyzed protein baits in McPhail traps. Here. ludens females to male pheromone (152). orange panels (reflecting maximally at 590 nm) attracted numerous mature A. The recent banning of the fumigant ethylene dibromide (EDB) by the US Environmental Pro­ tection Agency has finally prompted an upsurge in research aimed at identifying new postharvest treatments and monitoring tools (lures. when fly populations are small. is 3. chapote odor almost completely inhibited attraction of sexually active A. The odor of fermented yellow chapote (Sargentia greggii). have proven effective at capturing several Anastrepha species (e. Nevertheless. Entomol. only 3 1 . and is cumbersome to service (2). ludens (153). striata and A. Annu. 1 % were caught. Furthermore.annualreviews. were also effective at attracting sexually mature A. 51. A. the preferred native host of A. most importantly. and. which is baited with a mixture of protein (occasionally molasses or fermented fruit juices are also used) and water (2. 32. sphere attractiveness was increased further by placing calling males close to the balls (171). For personal use only. reflecting maximally within a narrow spectral region (Le. it works better in dry climates (50). 54. I only refer to relevant work in each of the areas that are currently being applied to manage Anastrepha populations. traps) . A. working in a mixed mango orchard with large numbers of A. Orange balls. it is very inefficient. and attempting to identify Anastrepha sexual pheromones in order to develop a pheromone trap.and overall management schemes. This trap. breaks easily. ludens. obliqua flies to McPhail traps in guava groves in Costa Rica (78). owing to space restrictions. 18. 500-590 nm). even though widely used. 117).166 ALUlA heavy reliance on postharvest treatments with fumigants. and A.6 times more attractive than yeast hydrolysate to hungry. 20 cm in diameter. by University of Guelph on 10/10/12.g.

Computer models suggest that the use of sterile pest releases and inundative release of parasitoids are more efficient together than either method alone . Of these. and Aceratoneuromyia indica have been imported and re­ leased in the US. Rev. suspensa flies (141). the impact that these release programs had on native fly p opulati ons was never adequately do cumented Examples of augmentative releases of parasitoids are rare with Anastrepha.annualreviews. Biological Control Both classical biological control and rep ea ted augmentative releases of mass-reared parasitoids have been used to suppress Anastrepha populations. unpublished data). These include efforts in Florida where thousands of D. longicaudata and D. 46. A. fraterculus in the US (86. Biosteres giffardi. by University of Guelph on 10/10/12. 89.a more critical selection of natural enemies to be released. Higher success rates in the future are. 128.D. Annu. 127. A. virgin A. Doryctobracon crawfordi. 122. longicaudata have caused significant reductions in wildA. Camacho. To date.several compounds have been isolated and their structures elucidated. ludens.Costa Rica.Mexico (48. Downloaded from www.39:155-178.contingent on prerelease studies. Such an approach has proven to be highly successful against Ceratitis capitata in Hawaii (195) under conditions similar to those encountered in Latin America.87).the current trend is to explore the possibility of joint sterile fly and parasitoid releases.Brazil. ludens (151). Enkerlin & T. iudens. 155. and a better understanding of parasitoid behavior and ecology. Unfortunately. H. J. unpublished data.and male-baited traps catch five to ten times more flies than food-baited traps in field trials using laboratory-reared. 187). and A. 1994.ANASTREPHA BIONOMICS AND MANAGEMENT 167 Identification of Anastrepha sexual pheromones has been a slow process that started in the 1970s and has been mostly restricted to two species: A. and A. Sivinski. suspensa. Entomol. Similar results have been reported for A. suspensa. Wong.Peru. For personal use only. Both females and males are attracted to calling males. 130). in my view. 154. ludens (77. 192). personal communication). crawfordi have become firmly established from the US to Argentina. Other efforts in Mexico and Costa Rica have also apparently been successful in controlling A. suspensa populations (19. ludens but have never been published (D.Y. Some of these compounds have been synthesized (22. . obliqua and A.Mexico. Sterile Insect Technique Even though release of sterile insects alone has proven effective at eradicating or substantially suppressing populations of A. fraterculus (47. 167).and Argentina for the control of A.T.and Peru (69). Classical biological control has been attempted repeatedly in various coun­ tries. B. Parasitoid species such as Diachasmimorpha longicaudata. suspensa and A. Ganaspis pelleranoi.

. Current studies are searching for toxicants that are more fly specific such as cyro_mazine ( 1 1 0) and avermectin Bl ( 1) and products that are less harmful to the environment such as borates (59). To date.annualreviews. and carambolas (72. 102. Since then. Rev. Because bait sprays have been shown to cause unacceptably high environmental damage. by University of Guelph on 10/10/12. /udens. and reductions in parasitization rates (56. Chemical Control Anastrepha flies are susceptible to almost all insecticides (2. A. Sivinski. Entomol. Also. as well as soil toxicants ( 168). and A. because sterile releases are most effective at low pest density. Such alternative disinfestation methods against A. and guava (164). guavas. parasitoids can be used to reduce fly popu­ lations prior to sterile fly releases.g. Legal Control Anastrepha is considered a major threat to the fruitgrowing industry in almost every country in the world. personal communication). grapefruit. rotenone. Also.168 ALUJA (20.39:155-178. fruit disinfestation procedures and establishment of fly-free zones) in the past 10 years. serpentina. For personal use only. surges of secondary pests. use of radiation (37). Another recent alternative is the establishment of fly-free zones. 179). acting poisons are preferred to prevent adult females from ovipositing after exposure to the poison. 16 1. 94). A. and cold-storage (71). the most widely used mechanism has been the use of bait sprays applied from the ground or air ( 102. disinfestation treatments must effect a mortality of 99. Quick­ Annu. but work is underway (J. In the early parts of the century. naturally occurring and inorganic insecticides such as nicotine. whereas the inundative releases of parasitoids are more efficient at high pest densities (20. no field tests reporting results of these new strategies applied to Anastrepha have been published. research is underway to find more acceptable alternatives. obliqua larvae have recently been developed: e. authorizing and strictly enforcing fruit disinfestation procedures. hot-water or hot-air treatments in mangoes. a flexible acoustical device was developed to detect feeding sounds of A. Downloaded from www. 166). suspensa larvae in such fruit as mango. were used. issuing of certificates and limited permits for use when fruit are moved from infested areas to places considered fly-free. Legal control encompasses quarantines. Thus a series of quarantine regulations limit the movement of fruit from infested areas to areas considered to be pest-free. orchard certification.9968% (probit 9) of eggs and larvae in the fruit (97). 1994. Several suspensa. 94). Generally. 158. . Most advances have been made in the two last areas (i. and establishment of fly-free zones. 163). and copper salts ( 13). Landolt et al (97) have proposed an alternative to this unrealistic value based on calculations of the probability of a mating pair surviving a shipment.e. 157). arsenic.

1 1). For personal use only. Our under­ standing of factors that regulate population numbers in the field and of how individuals can track down unpredictable and ephemeral resources is still only marginal. and most importantly.39:155-178. basic biology'and ecology. Comparative studies between monophagous and polyphagous. in unper­ turbed environments. and chemical ecology should be especially rewarding. Rev. and management. 1994. if at all possible. much more attention should be paid to the development of truly integrated (Le. For instance. (2. Ecuador. trap cropping. Downloaded from www. integrated . Other approaches currently under research are border trapping. One such substance is gibberellic acid. see also USDA-PPQ-APHIS Fly-free Area Regulations (unpublished infor­ mation).org by University of Guelph on 10/10/12. Alternative Methods A series of new approaches have been proposed recently. three areas of research warrant our immediate attention: taxonomy. countries could be divided up according to biogeographic charac­ teristics into large eradication zones. trap cropping. for a working example]. plant-growth regulators can cause the plant to produce more of a given resistance agent or sustain the innate resistance properties of the plant. behavior. develop economic thresholds tailored to meet both the stringent quality requirements of international fresh fruit markets and the less rigorous local markets. localized fly-free zones. life-history traits. Finally.annualreviews. Certification procedures are site specific and strictly enforced following a series of rules negotiated by both exporting and importing countries [150. habitat manipulation) and that are within the cultural and economic reach of the recipient (11). or primitive and derived. Mexico. DIRECTIONS OF FUTURE RESEARCH In my view. This effort might lead us away from species of economic importance but will undoubtedly add new depth to our under­ standing of basic patterns in Anastrepha biology and ecology. A thorough taxo­ nomic revision of the genus (including identification keys to immature stages and males) is long overdue.g.ANASTREPHA BIONOMICS AND MANAGEMENT 169 zones are currently operating in the US. species in areas such as demography. under natural conditions and. identify more efficient biological-control agents. In the area of Anastrepha management we need to develop better traps and lures. Entomol. integration of all available control methods and consideration of other pests and diseases that attack the same crop) and regional management schemes that are based on ecologically sound principles (e. As part of a regional management strategy. and habitat manipulation Annu. suspensa (74). which has been shown to reduce grapefruit susceptibility to attack by A. More emphasis needs to be placed on studying Anastrepha spp. and Brazil.

M Mangel. Liedo p. I. Dl 1 1-903537. Baker EW. In press Aluja M. 1994. Hendrichs J. Mexico. PhD . et al. Jacome I. Liedo P. 1993. 1983.170 ALUJA management zones. Baker AC. Review of concepts and recent developments. V Hernandez. Fla. and IFS (projects 05 1/93. Am. Plummer CC. McPhail by University of Guelph on 10/10/12. Studies on the Mex­ ican fruitfly known as Anastrepha fratercutus. RE Plant. obliqua. 122-33 Aluja M. See Ref. Guillen J. Behavi­ our of Anastrepha ludens. Quintero G. 7. New York: Springer Arrigoni EB. Entomol. Stone WE. 1993. A. 38:95100 ulacional de moscas-das-frutas (Dip­ tera: Tephritidae) em tres regioes do Estado de Sao Paulo. Celedonio H. Mane jo Integrado de las Moscas de la Fruta. 33-47 Aluja M. Brasil. 1993. respectively). A review of stud­ ies on the Mexican fruit fly and related Mexican species. 42a. Aluja M. Soc. ACKNOWLEDGMENTS I am most indebted to RJ Prokopy and RT Carde for their support. 12. Ann. Sao Paulo. 1945. Sherman M. de la Rosa G. lla. Natural host plant survey of the economically important fruit flies (Dip- Aluja M. 3. J. Ayora F. 12-48. I cannot list their names. 2nd ed. 70:329--38 tera: Tephritidae) of Chiapas. 1990. pp. Cabrera M. I sincerely thank all colleagues who provided reprints and copies of unpub­ Annu. J. and G Dieringer for extremely useful comments on a previous version of the manuscript. Albrecht CP. Mexico. Guillen J. En­ tomol. 80: 344--47 Aluja M. 1993. Fla. Cabrera M. Entomol. Appl. 2. lished manuscripts. Guillen J. and Cl 174 1-1. pp. 6. The study of movement in tephritid flies. Aluja M. Brasil. 1984. A Norrbom. Ann. 13. Lethal and sublethal effects of Avermectin B I on three fruit fly species (Diptera: Tephritidae). Cabrera M. Entomol. Sao Paulo. et al. 1986. Unusual calling be­ havior of Anastrepha robusta flies (Dip­ tera: Tephritidae) in nature. 162 pp. I acknowledge support from Programa MOSCAMED (DGSV-SARH). 8. Univ. Entomot. SEP. ser pentina (Diptera: Tephritidae) on a wild mango tree (Mangif era indica) harbouring three McPhail traps. In press Aluja M. Fruit infesting tephritids (Dipt. Mexico City: Trillas. 531:1-155 14.annualreviews. but because of space restrictions. Fruit Flies: Biology and Management. pp. Birke B. Econ. and zones where any control action is technically and economically unfeasible (see 1 1).39:155-178. Dinamica pop­ thesis. Lozada N. Future per­ spectives on integrated management of fruit flies in Mexico. 1989. Literature Cited 1. Entomol. USDA Misc. P Liedo. Publ. 1987. Am. Hendrichs J. Operations and Systems Analysis in Fruit Fly Management. eds. Downloaded from www. 252 pp. Cabrera M. JR Carey. Econ . I thank P Greany. Rev. Ber­ lin/New York/Tokyo: Springer Aluja M. Entomol. 5. de la Rosa G. 11a. Rios E. Lekking behavior and male territo­ riality. 4. ed. Birke A. 1944. Rios E. Basic patterns of behavior in wild Anastrepha striata (Diptera: Tephritidae) flies under field­ cage conditions. I. Liedo P. DGICSA-902467. 10:309-18 Aluja M. Aluja M. In Pest Control: 9.: Tephritidae) and associated parasitoids in Chiapas. obliqua under seminatural conditions. For personal use only. See Ref. In press II. andA . Soc. 1987. CONACyT. J Sivinski. Entomophaga 35:39-48 10. Insect Sci. 1993. 1993. General behavior and interactions between Anastrepha ludens and A . Habitat use by Anaslrepha obliqua flies (Diptera: Tephritidae) in a mixed mango (Mangif era indica) and tropical plum (Spondias purpurea) orchard.

Bras. Appl. Entomol. Probability of detecting Caribbean fruit fly. 35:67984 Bressan S. Webb JC. Dispersal and orientation of sterile Ceratitis capitata and An­ astrepha ludens (Tephritidae) in Chiapas . 22. 31. Entomal. Entomo/ . Rev. da Costa-Teles M. Fla . Webb Ie. 1988. Enlomol. Anastre pha suspensa (Loew) (Diptera: Tephritidae): territorial fights and signalling stimu­ lation. Ramos MRK. Carvajal SSR. Profundidae de pupaltao de Anastrepha obliqua (Macquart. Anastrephin and epianastrephin. The cytotaxonomy of the larvae of some Mexican fruit flies in the genus Anastrepha (Diptera: Tephritidae). An. Entomol. 1982. Longevidad e curva de sobrevivencia de tres especies do genero Anastre pha Schiner. . Chan AST. 20. Soc. 1 99 1 . Ann. Vanderbilt DP. Wasps sting flies. Chan AST. 76: 678-82 Bush GL. Theor. Environ. song parameters. Jimeno-Zavala MA. J. 1983. Downloaded from www. Guillen J. Berrigan DA. Toledo J. 41 :75-79 Bressan S. 1972. Appl.39:155-178. 17. 20:1 8-26 Burditt AK. 1 12:410-2 1 Baker PS. 17: 493--5 1 8 Battiste MA. and mating success in Car­ ibbean fruit flies Anasrrepha suspensa (Loew). Rev. 66:330-44 Burk T. (Diptera: Tephritidae) liberados em apenas urn ponto do pomar. 1983. The ecology of fruit flies. 36. Bioi.1 0 1 Bustos ME. . Invest. 30. Brasil 19:471-79 Bressan S. Pictorial Key to Fruit Fly Larvae o f the Family Tephritidae. 1 1 2:263--73 Baker PS. Saleh SM. da Costa-Teles M. Brasil. Brasil. 1 :30-33 Braga MAS. BioI. 1 99 1 . Anastrepha suspensa (Loew) (Diptera: Tephritidae). Econ. Tephritidae) em laborat6rio. 19. 23. Ecol. J. Las moseas de Jrutas del genero Anastrepha Schiner. See Ref. 171 16. Anastrepha ludens. 79:409-16 Calza R . 35 . En­ romol. Fla. Baker P S . Appetitive dispersal of sterile fruit flies: aspects of the methOdology and analysis of trapping studies. (Diptera: Tephritidae) em condi"oes naturais . Anastrepha sus­ pensa (Loew) (Diptera: Tephritidae) populations with McPhail traps . Male-male interactions in Caribbean fruit flies. Research 87 3:12-13 Barclay HI. Entomol. Nation JL. da Costa-Teles M. 65:367-73 Burk T. Levantamento de "moscas-das-frutas" do genero An­ astrepha em vanos municipios de Sao Paulo. Arq. 67:542-48 Burk T. Schroeder WG. 77: 1 98-201 Calkins CO. Rev. Sao Paulo 55:55-60 Caraballo J. 1972. Entomol. 29. Carrasco H. 25. Appl. An. 28. Inst. 1962. 1 987. Lopes FD . Entomol. 1 99 1 . Behavioral ecology of mating in the Caribbean fruit fly. Chambers DL. Psyche 69:87. 60-40. Effect of male size on calling propensity. 1994. 341-44 Calkins CO. 1 98 1 . Soc . San Salvador: OIRSA . 27. 37. da Costa-Teles M . Suplicy N Ir. 47:73-80 Blanchard EE. 1969. 1868 Annu. Mexico . Popul. Celedonio H. Fla. 1993. Trapping for control of the Mexican fruit fly in mango and citrus groves. 34. pp. For personal use only. 38. 15:28 1-342 Boush OM. Exp. 1 96 1 . Strekowski L . 1 98 1 . 35:685-90 Bressan S. Visnick M. 24:26 1 1-14 Berg GH 1979. 24. Entomol. Entomol. 39. Entomol. Influencia das cores e formas das annadilhas na captura de Anastrepha spp. 36 pp. Bacteria associated with the Caribbean fruit fly. 1988. 1988. King RW. Entomol. 1984.ANASTREPHA BIONOMICS AND MANAGEMENT 15.) (Diptera: Trypetidae). 33. 1 99 1 . 26. Rev. 23:2738 Balock JW. 1983. Bioi. 1990. Recaptura de adultos marcados d e An­ astrepha spp. Zucoloto SF. 1987. 1 99 1 . Am. Agric. J. 21 . Guidelines for a sterile release pro­ gramme. Entomol. J. Es­ tudios sobre a melhor concentraltao de aminOlicidos para moscas adultas de (Diptera: obliqua Anastrepha Tephritidae). Fla . £Con .annualreviews. Brasil. Temporal and seasonal differences in movement of Caribbean fruit fly larvae in grape­ fruit and the relationship to detection by acoustics. 1 868 (Diptera. J. Entomol. 1 835) (Diptera: Tephritidae) em tres substratos. Enkerlin WR. Rev. Carey JR. 18. Annu. Models for pest control: complementary effects of pe­ riodic releases of sterile pests and parasitoids. novel lactone components from the sex phermonone blend of male Caribbean and Mexican fruit flies. En­ tomol. 32 : 7689 Bateman MA. lat. 40. 1984. Irradiation as a quarantine treatment for Mexican mangoes. Baranowski RM. Age and host effects on clutch size in the Mexican fruit fly. Anastrepha suspensa (Loew) (Diptera: Tephritidae) McPhail traps for survey and detection. App/. Tetrahedron Lett. Reyes J . l l a. Rev. Soc. Entomol. Chan AST. 1 by University of Guelph on 10/10/12. 62:53-57 Baranowski RM. Quanti­ fication of tephritid fruit fly dispersal. Raga A. Especies Argen­ tinas del genero Anastrepha Schiner (sens. 32. 41.

Fla . En­ tomol. Cient(jica 17: 1 89-93 Ferro MIT. Biologia e comportamento de Anastrepha grandis 44 62. Proc. 47. Costa-Lima A. PhD thesis. Calkins CO. 5 : 1 7 1-92 Chuman T. Entomol. USDA Agric. 1930. Ga. Enkerlin WR . 1 984. ed. Rev. Villalobos R. Rev. Wharton RA. Rotterdam: Balkema Celedonio-Hurtado H. Mexico durante 1 966. 1. Suda DY. Soc. 44. Tephritid fruit fly trapping: liquid baits in high and low rainfall climates. Nakagawa S. Suspensolide. Anastrepha ludens (Loew) . hydrolized protein. Carey I. Bi­ ology of fruit flies.P. 1 934. 7 1 :762-63 Cytrynowicz M. C. 63. Influencia da nutril<ao proteica no desenvolvimento da gHindula salivar de machos de Anastrepha obliqua Mcqu­ art. 1 990. esteriles y silvestres. Entomol. Fit6 filo 19:43- 42. Bull. to colored rectangles and spheres. Wash. 1933. Carroll LE. ritidae). 1984. 1984. borax. et al. Tumlinson JH. Entomol. 52. (D iptera : Tephritidae) in Guate­ mala. Agric. Efeito do balan\<o de protefna e carboidrato na dieta de Anastrepha obliqua Mcqu­ art. Davis IC. Trap features that promote cap­ ture of the Caribbean fruit fly. 1 :236-48 55. Sobre insectos que vivem em maracujas (Passiflora sp. pp. Reyes J. 1988. Brasil. Anastrepha ludens Loew (Diptera: Tephritidae): an inte­ grated behavioral and ecological study. Ehler LE. 1932. de Souza HML. Envi­ ron. 49. citrus and walnut. 82: 201-14 Cavalloro R. 1982.L. Central de Venezu­ ela. The biology of dacinae fruit flies. 1 1 : 1 202-10 da Silva-Gomes J. Petropolis 2:302-10 Fletcher B S . Morgante IS. ludens. 1 89-200 56. N. Mor­ phology of the immature stages of Anastrepha ludens (Diptera: Teph­ ritidae). Univ. ITESM­ Monterrey. 480: 1-545 Cons-Duarte M. Rev . Univ. a new macrolide component of male Caribbean fruit fly [Anastrepha suspensa (Loew)] vola­ tiles. Zucoloto FS. Liberaciones de mosca Mexicana de la fruta (A nastrepha ludens) esteril en Baja California. Soc. 46. 1982. Nota taxon6mica e biol6gica sobre Anastrepha grandis Macq. (Diptera: Tephritidae) . Nasca AI . Knapp EM. 43. USDA Tech . Especies d e braconidae (Hyme­ noptera: Ichneumonidae) parasitoides de moscas de las frutas (Diptera: Tephritidae) colectados en la provincia de Tucuman (Argentina) . Tephritidae). Maracay. Trujillo­ Garcia P. Rev.39:155-178. Entomol. Aluja M. 1 1 0 pp. Annu. 53. En­ tomol. The eggs of four species of fruit flies of the genus Anastrepha. UK. 1988. Berrigan D. See Ref. En­ tomol. 35 : 1 84-9 1 Enkerlin WR .. Zucoloto FS. Chambers DL. Demography of A nastrepha ludens. Sivinski I. 1987. 29:6561-64 Clausen. Effect of malathion-bait sprays on biological control of insect pests of olive. Fruit Flies o j Economic Importance. Southampton. Heath RR. Inst. Tephritidae). 1 8 1 pp. 210 pp. 1960. 1987. For personal use only. 76 pp. (Maequart. Parasitism of fruit flies Ceratitis capitata and Anastrepha spp. 1978. Endicott PC. Tech. Entomol. Entomol. 353-58 Eskafi FM. 1989. 1989. Guillen I. 1846) (Diptera: Teph­ 65. Darby HR. 60.). Oswaldo Cruz 23: 15962 Cunningham RT. An­ astrepha Jraterculus. ob/iqua and A . 7 1 : 1 1 1-20 Chan AST. Mexico. Foote RE. Studies 66. 42a. and Mediterra­ nean fruit flies. 1978. 1989. 1980. Entomophaga 35:355-62 Fernandez-de-Araoz D. 51. on the Mexican fruit fly . 1966. Orientaci6n y dispersion de poblaciones de la mosca Mexicana de la Jruta (Anastrepha en el municipio de Allende. 64. Downloaded from www. . 58. Hilgardia 52:1-47 57 . Mating and terri­ toriality in wild Anastrepha suspensa (Diptera: Tephritidae) in field cages. Venezuela. Liedo P. MS thesis. Eeon. and water to control Anastrepha fruit flies. Annu. A. en el per(odo de septiembre de 1985 a agosto de 1986. Christenson LE. 1 7 . Introduced para­ sites and predators of arthropod pests and weeds: a world review. Urago T. 1994. 1835 (Diptera. Mem. 1835 (Diptera. Dodson GN. Loew). Ann.172 ALUJA (Di ptera: Tephritidae) de Venezuela. CIRPON 2:37-46 Ferro MIT. PhD thesis. 59. 32: 1 1 5-44 Foote RH. 50. 444: 1-20 54. 1983. Soc. Entomol. Fruit fly genera south of the United States (Diptera: 45 . Ceratitis capitata. Cient(jica 17: 1-5 Fischer CR. Univ. Sao Pauio. Annu. Tetrahedron Lett.annualreviews. 1. serpentina (Diptera: Tephritidae) in Mexico. 199 1 . Entomol. Am. l l a. Visual responses of South American fruit flies. 1993. 1. MS by University of Guelph on 10/10/12. Agee HR. Emmart EW. 1989. Bull. Patton WP. Use of a mixture of boric acid. 61. 48.

Sharp JL. Harris DL. Consum. Entomol. Dis­ tribuci6n y Plantas Hospederas. 1989. Rev. Mexico: Inst. Ithaca: Cor­ nell Univ. Cooley SS. 1993. . Handbook o f the Fruit Flies o f America North o f Mexico. Increasing effec­ tiveness of visual traps for the Carib­ bean fruit fly . Vargas C. Una sustancia nat­ ural en la captura de moscas de la 173 67. f ertility. Calif. Sivinski J. Swe­ den.39:155-178. 30: 1 40--5 0 Jimenez-Jimenez E. Jara B. 162 pp. PhD thesis. Econ. J. 81. 1994. Res. J. 70. Mazor M. Rev. 78. Uppsala. Appl. J. D .). Soc. Fla. Downloaded from www. Entomol. McDonald RE. Aluja M. Blondheim S . 1 : 1-30 Holler TC. MS thesis. Characters of the larvae and pupae of certain fruit flies. Entomol. Suarez A. 75. Hernandez-Ortiz V. EI gusano de la frota (In­ strypetas ludens I . Univ. Press Galun R.annualreviews. Improvements in efficacy of gibberelic acid treatments in reducing susceptibility of grapefruit to attack by Caribbean fruit fly. Norrbom AL. Entomol. Cold-stor­ age quarantine treatment for carambolas infested with the Caribbean fruit fly (Diptera: Tephritidae) . Development of at­ tractants for monitoring Caribbean fruit flies. 79 pp. 1993. Mex. Folia Entomo!. B . Blanc FL. Tejada LO. Fla. Landolt PJ. Berkeley. 1990. 82. 1990. Rio Grande Valley Hortie. Epsky NO. Entomol. Exp. 84. 261 pp. Chambers DL.ANASTREPHA BIONOMICS AND MANAGEMENT Tephritidae). 1956. 1978. 73. 83 . J. Schroeder WJ. Uptake of plant surface leachates by apple maggot flies. Econ . See Ref. Anastrepha interrupta (Schoep­ fia fruit fly) (Diptera: Tephritidae). 14:726-32 Gonzalez J. 1 99 1 . Comparison of aggregation and feeding responses by normal and irra­ diated fruit flies. In press Herrera AL. 68. The guava fruit fly. Annu. Las moscas de la fruta y sus enemigos naturales. Population fluctuations of economic species of Anastrepha (Diptera: Tephritidae) re­ lated to mango fruiting phenology in Costa Rica. Entomol. 76: In press Hedstrom 1. 74:570-80 Greene CT. Anastrepha fraterculus (Wied. Bull. 1 952 . 1929. Dept. Xalapa. Garcia R. HedstrOm I. Ecologfa-Soc. 1 12 pp. Mex. 72. 197 1 . 23:20-25 Greany PO. Fluctuations in numbers and the significance of the sex ratio of the Mexican fruit fly. pp. 74:98-105 Jiron LF . Comision Parasitol. En­ tomol. 77. 173-75 Heppner JB. 1993. fruta del genero Anastrepha Schiner (Diptera: Tephritidae). Entomol. 1979. Mexzon RG. Fla . 1984. No. Agric. BioI. Sharp JL. 1986. Ceratitis capitata and Anastrepha suspensa (Diptera: Teph­ ritidae) . Anastrepha ludens caught in McPhail traps. and longevity o f three tephritid species. 79 . Rev. Folia Entomol. 87. Watts. USDA Tech. 4 1 :4960 Gould WP. 69. 91 . Lachman A . . Foote RH. 38:489-504 Hagen KS. 198 1 . Bal. Peru. Fla. Heath RR. 43 pp. Uppsala Univ. 1985. El Genero Anastrepha Schiner en Mexico (Dip­ tera: Tephritidae): Taxonom( by University of Guelph on 10/10/12. 1900. PhD thesis. by use of flu­ orescent colors. 83:458-60 Gould WP. 1993. Florida. 1 600 (SEA). Release of sterile Mexican fruit flies for control of feral populations in the Rio Grande Valley of Texas and Mexico. 1 4:66-86 GonzaIez-Hernadez A. Agee HR. Fluctuacion de la poblacion de Anastrepha ludens (Loew) y de sus enemigos naturales en Sargentia greggii S . Efficacy of sterile release of Anastrepha sus­ pensa adults against wild populations. 80. Hendrichs J. Agric . 36:269-72 Hedstrom 1. 1992. Hendrichs J. in seasonal and non-seasonal neotropical f orest environments . 1992. 6. Entomol. See Ref. Gothilf S . 37: 1 1 3-27 Holler TC. Agric.). 327-33 Houston WWK. Parasitoid . For personal use only. Burditt AK. Estudios sobre l a aplicacion de la tecnica de machos esteriles en el control de la mosca Sudamericana de la fruta. 1988. Davidson JL. Hot-water immersion quarantine treatment for guavas infested with Caribbean fruit fly (Diptera: Tephritidae). Entomo/. Gainesville. de la Barreda A. 74. Larvae of fruit flies. Rangel AF. 85 . 1 99 1 . Lista preliminar de especies del genero neotropical Anastrepha (Diptera: Tephritidae) con nolas sobre su distribucion y plantas hospederas. Servo En­ tomol. 327: 1-2 Hernandez-Ortiz V. Ha. Anastrepha striata Schiner (Teph­ ritidae). 86. Univ. Prokopy RI. Trop. Anastrepha suspensa (Loew) (Diptera: Tephritidae). 90. Environ. Fit6 filo 9:4-1 1 Jiron LF. 71. 89. Exp. Appl. 85: 1 235-39 Greany PO. 88. Sharp JL. 1 99 1 . Sexual selection in wild and sterile Caribbean fruit flies. 76. 1 1 a. Circ. Shaw PE . 1993. Entomo/. Mex. Anastrepha suspensa (Diptera: Tephritidae). pp. Influence o f adult nutrition upon f ecundity. Anastrepha suspensa (Diptera: Tephritidae).

Morgante IS. 63: 135-42 L6pez F. Browning HW. Size specific demography of three species of Anastrepha fruit flies. Entomol. Prokopy RJ. Distribucion ecologica de especies del genero Anastrepha Schiner en el noroeste Peruano. 7 1 : 130-37 93. Bliss CI. Econ. Entomo!. Immature stages of the Caribbean fruit fly. 1 1 3. 109. Norrbom AL. Distribution and activities of Anastrepha fraterculus (Diptera: Tephritidae) flies on host and nonhost trees. Mex. Genet. 1992. Rev. 99. Control of the Mexican fruit fly by bait sprays concentrated at discrete locations. Ann. 5:263-78 Malavasi A. 1 969. 1 988. 1989. Fla. Am. Korytkowski C.39:155-178. 1 I :32-70 96. 1868 en el noroeste Peruano. by University of Guelph on 10/10/12. 1 992. 1 933. Pop­ ulation genetics of Anastrepha fratercu!us (Diptera: Tephritidae) in different hosts: genetic differentiation and heterozygosity. Fla. Chambers DL. Environ. Es­ pecies del genero Anastrepha Schiner. J. 1992. Rev. Environ. 62: 1 255-57 L6pez F. Malavasi A. Kitto GB 1 983. 74:1 89-96 Leyva JL. Ill. 103. Steiner LF. Entomo!. 20: 1 I6065 Liedo P. Kamasaki H. Observa­ tions on the Mexican fruit fly and some related species in Cuemavaca. 1 I2. 1 5 :33-50 McPhail M. Solferini VN. Trop. 1 983. Carey J. A preliminary list of the fruit flies of the genus Anastrepha (Diptera: Tephritidae) in Costa Rica. 1986. 101 . J. Chambers DL. Biochem. Practical implications for regu­ lating insect populations by natural enemies.annualreviews. Holbrook FR. 97. Baranowski RM. Gilstrap FE. hymenopterans of Costa Rica: geo­ graphical distribution of the species associated with fruit flies (Diptera: Tephritidae). 36:55-65 Martinez AJ. Exp . Berry NO. Econ. J. 1 98 1 . Agron. Korytkowski C. 102. Chew V. 108. Folia Entomol. obliqua (Diptera: Tephritidae): ev­ idence for gene duplication. See Ref. Genet. Entomo!. Entomophaga 34:53-60 92. 1 969. Guillen J. Knipling EF. Rev. Rev. Bras. 1936. 105. Morgante JS. Alternative to the use of Probit 9 mortality as a criterion for quarantine (Diptera: treatments of fruit fly TephritidaeHnfested fruit. Entomol. l iS . Morgante JS. Godoy F. 42a. Development of Anastrepha ludens (Diptera: Tephritidae) in several host fruit. Genetical and biochemical com­ parisons of alcohol dehydrogenase iso­ zymes from Anastrepha fratercu!us and A . 64: 1541-43 Malavasi A. 1994. 1 979. Ge­ netic variation in natural populations of Anastrepha (Diptera: Tephritidae). 1988. Landolt PJ. 107. 1 14. Econ. 76:28692 Malo E. Entomo!. 104. Folia En­ tomol. 77:285-87 Lawrence PO. southern Mexico. Entomol. Malavasi A. 1968. Sanchez­ Riviello M. 62:214-19 Leyva JL. Morgante JS. A new yeast hydrolysate-borax bait for trapping the Caribbean fruit fly. USDA Agric. J. Soc. 24: 1 3-24 Matioli SR. Celedonio H. 106. Soto-Manatiu J. 1 983. Econ. Morgante JS. 73: 125-40 Martinez AJ. Effect of cyromazine on the oviposition of Mexican fruit fly (Diptera: Tephritidae) in the laboratory. App!. Genetica 60:207II Malavasi A. 1 982. Valenzuela J. Downloaded from www. An immunological approach to the phylogeny of the Tephritidae. Morgante JS. Moreno DS 1991. 1 984. 1991. Handbook 693: 1-337 95. pp. 29:405-1 1 McPhail M . Entomo!. Obser­ vations on Anastrepha palLens (Coq. For personal use only. 1 97 1 . 98. 10:275-78 ship to host availability. J. Econ. 1 986. Baker PS. Entomol. Mex. Peru.174 ALUJA tomol. Temperatura umbral y unidades calor requeridas por los estados inmaduros de Anastrepha ludens (Loew) (Diptera: Tephritidae). Entomo!. Genet. Adult and larval population flUctuations of Anastrepha fratercu!us and its relation- 12:7 1-95 1 I0. Anastrepha suspensa. J. Frias D.) reared from wild fruits in the lower Rio Grande Valley of Texas during the spring of 1932. Principles of insect parasitism analyzed from new perspec­ tives. . 100. Jiron LF. Entomol. Entomol. Ojeda D. 84:1 540-43 Mason U. 203-1 1 94. Brasil. Rev. 82:1 39-42 Matioli SR. Response of Caribbean fruit fly (Dip­ tera: Tephritidae) to modified McPhail and Jackson traps: effects of trapping duration and popUlation density. 1987. Evolutionary trends of alcohol dehydrogenase isozymes in some species of tephritid flies. Econ . In­ fluencia de los factores meteorol6gicos sobre la fluctuaci6n poblacional de Anastrepha obliqua McQuart (Diptera: Tephritidae) en mango. En­ Annu. Ojeda D. Entomol. The abundance of species of An­ astrepha (Diptera: Tephritidae) in the coffee producing area of coastal Chiapas. Entomo!.

. Downloaded from www. 1989. Insect Morphol. 125. . 1 34. 1 17 . Revision of the schausi group of Anstrepha Schiner (Diptera: Tephritidae). The species of Anastrepha (Diptera: Tephritidae) with a grandis-type wing pattern. Temporal dynamics of host-marking in the trop- 127. 8:24 1-47 Morgante JS . and partial chemical characterization. 1 33-47 Mori K. Parasitos de "Mosca de la Fruta" establecidos en algunas zonas de Tucuman. Entomol. Evolutionary pat­ terns in specialist and generalist species of Anaslrepha. Efeitos das misturas de alguns al­ imentos na produ<. 95:52-58 Norrbom AL. Brazzel JR. Colombia. Malavasi A. Environ.annualreviews. Am. 1992. Ceratitis. Embryol. Am. l 1 a. 128. 93: 1 0 1-24 Norrbom AL. and Ceratitis capitata (Diptera: Tephritidae) in Brazil. 1 99 1 . pp. Econ. 1988. Duarte AL. Dacus and Rhagoletis (Diptera: Tephritidae). State Univ . No. with a discussion of the terminology of the female terminalia in the tephritoidea. Entomol. 1992. The sex pheromone blend of Caribbean fruit fly males: isolation. 135. Two new species of Anastrepha (Diptera: Tephritidae) with atypical wing patterns. Sex-specific glands in tephritid fruit flies of the genera Anastrepha. Fruits 45:629--3 1 Nasca AJ. Rev. 4:27-30 129. Rivera P. 136. Hernandez F. Dinamica poblacio­ ysis and taxonomy o f the cryptostrepha. Liebigs Ann. 120. 1 38. Norrbom AL. Courtship behavior and evidence for a sex attractant in 175 1 16 . Biochemical systematics and ev­ olutionary relationships of neotropical 73:622-30 Morgante JS. Chem. J. Nakazono Y. Ann. Nation JL. Prokopy RJ. nal del comple jo constituido por las moscas de las /rutas Anastrepha striata y Anastrepha fraterculus en el media ecol6gico del sur de Santander. 65:1364-67 Nation JL. Evaluation of six different traps for detecting the Mexican fruit fly . dacif ormis. Soc. Agron. Bush GL. 1983. 1993. 1993. 66:234-41 Morgante JS. 1972. Papaj DR. Selivon D. 1985. 1975. Soc. 1994. 1993. J. Pesq. Nguyen R. Brasil. . Wash. Rev. Soc. Kim KC. 1985. II. 6:595-98 Montero CAB. The taxonomy and zoogeography of the genus Anastrepha (Diptera: Teph­ ritidae). 1988 : 1 67-74 Murillo T. Univ. Florida. Seasonal occurrence of An­ astrepha suspensa (Diptera: Teph­ ritidae) in Indian River County . En­ tomol. 75 pp. Chem. Jiron LF. Nation JL. 16:553-72 1 3 1 . Ge­ netic variability in populations of the South American fruit fly Anastrepha Jraterculus (Tephritidae). Soc. Anastrepha. J. Wash. USDA-APHIS Misc. Entomol. and schausi spe­ cies groups o f Anastrepha Schiner (Dip­ tera: Tephritidae). 1 990. Solferini VN. Sanchez-Salas JAS. Norrbom AL. Anastrepha suspensa. 133. 1989. 1 23 . 85:456-60 Nascimento AS. Malavasi A. Ann. Am. For personal use only. Brasil. 1 14 pp. 122. Malavasi A. Flutua«. Kim KC. 1 18. Aluja M. USDA Circ. Soc. Entomo!. ob­ liqua. 17:969--80 Nation JL. Brasil. Proc. Poucher C. populacional. Rev. 3A: 1 5-26 Norrbom AL. 139. Entomol. 1984-1987. Bucaramanga. 85: 8 1 3-20 1 32. Anastrepha obliqua (Diptera: Tephritidae). A list of the reported host plants of the species of Anastrepha (Diptera: Tephritidae). 1 990. Undustrial de Santander. 1988. Penn. 10:3 1-43 Nascimento AS. Dinamica pop­ ulacional das moscas-das-frutas do genero Anastrepha (Diptera: Tep­ hritidae) no Reconcavo Baiano. MS thesis. 1 5:327-3 1 Morgante JS. En­ tomal. Norristown. Foote de ovulos em Anastrepha obliqua (Diptera: Teph­ ritidae). Ecol. Synthesis of lactone components of the phero­ mone of Anastrepha suspensa. Noroeste Argent. Zucoloto FS. Southwest. Publ. Rev. 10: 1 21-29 130.39:155-178. 81-52. robusta. Anastrepha suspensa (Diptera: Tephritidae). En­ tomol. Matioli SR. Malavasi A. Phylogenetic anal­ male Caribbean fruit fly. See Ref. biological activity. 137. Indigenous microflora of the West Indies fruit fly. Proc. 1982. 8 1 : 1 64-73 Norrbom AL. sus­ pensolide. Entomol. Malavasi A. and the enantiometers of anastrephin and epianastrephin. Biology of phero­ mone release by male Caribbean fruit flies. Zool. 255: 1-24 Message CM. Ann. Econ. 1973. 1 54a. 1 990. Genet. 1980. Entomol. Hot-water im­ mersion treatment for mangoes infested with Anastrepha !raterculus. Zucchi RA. Annu. See Ref. 1 19 . 124. Int. A . Agropecu. 1 98 1 .ANASTREPHA BIONOMICS AND MANAGEMENT Mexico in 1928 and 1929. Entomol. PhD thesis . 121. by University of Guelph on 10/10/12. Olarte WE. Mating behavior of wild Anastrepha f raterculus (Diptera: Tephritidae) on a caged host tree. 126. 355 pp. Morgante JS. 1980. Colom­ bia. Fla. Morgante JS.

14 6 . Amsterdam: Sharp JL. 3: 79-86 Perdomo AJ. Entomol. Tor­ res-Rivera CN. pheromones of the Mexican fruit fly. Colomb. 1977. 79:706-8 162. to com­ binations of fermenting fruit odor and male-produced pheromone in laboratory bioassays. Ecolog(a and male-baited traps under field con­ ditions. 160. 407-13 Robacker DC. Chem. New host plant and parasitoid record for Anastrepha alveata Stone (Diptera: Tephritidae). 8:763-7 1 Prokopy RJ. Soc. 72:41-49 Riherd C. 92 pp. J. 1986. 1985. a native host of the Mexican fruit fly. Hooper G. Hallman GJ. Belloti AC. 1990. Roitberg BD. 159. proteins. 85: 168-71 164. Entomol. See Ref. Aluja M . Entomol. Franqui RA. Econ. 1980. Oviposition deterring pher­ omone in Anastrepha suspensa . J. Oviposition deterring pher­ omone in Anastrepha Jratereulus flies. Ecol. 1976.S)-(-)-epianastrephin: male produced 140. 147. Am. En­ tomol. Entomol. 158. USDA Tech. eds. Proc. Entomol. 1990. J. 19. Monk JW. 1976. J. Entomol. Wash. Sharp JL. Battiste MA. 1 984. Ecol. 1993. Eeon. Kovaleski A. Entomol. Entomol. Physiol.annualreviews. Ann. 1984. Ecol. Comparison of flight ability of wild-type and laboratory­ reared Caribbean fruit flies on a flight mill. Am. Univ. Appl. Garcia JA. d a Silva MT.1 76 ALUJA ical tephritid fly. Soc. In press 153. (Z. 1994. 1988. 73:631-33 1 6 1 . McPhail M. 18:223-44 Rubio REP. Santiago LR. Natural Enemies and Control. 152. 156. Phillips VT 1 946. Nation JL. Thalman RK. Hart WG. Baranowski RM. Malavasi A. Hot water dip treatments to destroy Anastrepha ob­ liqua larvae (Diptera: Tephritidae) in mangoes from Puerto Rico. 142. The biology and identification of trypetid larvae (Dip­ tera: Tephritidae).Z)-3. 1990. McFaden MW. Rev. J. Isolation and identification of bacteria in the digestive tract of the Mexican fruit fly.6-nonadienol and (S. Robinson AS. Anastrepha suspensa. to components of male-produced sex pheromone. Puerto Rico 74:441-47 Sharp JL. Downloaded from www. 141. 1. Robacker DC. pp. 1992. Environ. Attraction of a laboratory strain of Anastrepha ludens (Diptera: Tephritidae) to the odor of fermented chapote fruit and to pheromones in laboratory experiments. l l a. Environ. Sharp IL. 145. Envi­ ron . 1982. Mexico. Agric. Re­ sponses of laboratory-strain Mexican fruit flies. Webb IC. Citrus production areas maintained free of Caribbean fruit fly for export certification. Univ. Xalapa. ed. Anastrepha ludens. Ga. Attraction of female and male Caribbean fruit flies to food-baited 154. 12: 1-161 Piedra E. Hot-water treatment for control of Anastrepha suspensa (Diptera: Tephritidae) in mangoes. Horti Sui 1 : 10-1 1 Segarra-Carmona AE. pp. For personal use only. Hot-air quarantine treatment for carambolas infested with Caribbean fruit fly (Dip­ tera: Tephritidae). Flexible acoustical device to detect feeding sounds of (Diptera: Caribbean fruit fly . Flight propensity of Fruit Flies: Their Biology. RamIrez-Ramos LV. Comparison of volatiles emitted by male Caribbean and Mexican fruit flies. Entomol. Chambers DL. Elsevier Rocca JR. In­ seticidas para controle da mosca-das­ frutas. Morgante IS. 149. In Fruit Flies. 1 4:17 15-26 Robacker DC. Zuiiiga A . Sharp JL. Strekowski L . Anastrepha ludens (Diptera: Tephritidae). Zuiiiga A. 1. Anastrepha ludens. 1 990. 1 9:403-8 Robacker DC. Rev. Am. En­ tomol. 157. 6:463-65 Prokopy RJ. 1989. Soc.86. 295-30 1 . Exp. Bull. 39: 1 03-8 155 . Behavioral re­ sponses of female Mexican fruit flies.39:155-178. 1 6:2027-38 de las moscas de la Jruta del genero Anastrepha (Diptera: Tephritidae) en Llano Grande y Monte Blanco. and amino acids by Anastrepha sus­ pensa (Loew) (Diptera: Tephritidae) in the laboratory. Soc. 1 941 . Estudios sobre las moscas del tallo y fruto de yuca: Anastrepha pickeli y Anastrepha manihoti. Amsterdam: Elsevier Prokopy RJ. For­ aging behavior of true fruit flies. Entomol. 95: 1 27 Plummer CC. Entomol. Veracruzana. Con­ siderations on the reproductive behavior of Anastrepha pseudoparallela Loew 1 873 (Diptera: Tephritidae). Chem. BS thesis'. 59: 1 0 1 5-16 Salles LAB . Piedra E . Con­ sumption of carbohydrates. 1992. Garcia JA. 775 : 1-12 Polloni YJ . Sci. J. Hart WG. Anastrepha ludens. Vera­ cruz. 1977. 1. 5 : 1 208-10 Peiia ]E. Masuda S. 1 3 : 768-73 163. The yellow chapote. Mem. Greany PD. 154a. Chambers DL. 1 993. AP Economopoulos. Eeal. 1966. by University of Guelph on 10/10/12. Entomol. Chem. Entomol. 1988. Chem. Environ . Nation JL. 1993. 1 1 :255-58 Sharp JL. 150. 143. 144. Econ. 15 1 . Annu. (Z)3-Nonenol.

1 3:201-8 Soto-Manatiu J. Chambers DL. Food Chern. Entomol. Webb JC. Soc. Soc. 1987. Steck GJ. Sao Paulo. 1983. 54:666-68 Shaw JG. Publ. Wash. Isolation and identification of novel lactones from male Mexican fruit flies. Econ. Anastrepha suspensa. Pictorial key to species of the genus Anastrepha (Dip­ tera: Tephritidae). Entomol. Entornol. Bull. Steck GJ. Environ. 1 989. grandis (Diptera: Tephritidae). Ann. 1 990. J. Anastrepha (Diptera: Tephritidae) associated with 177 mango 165. et al. 1977. 1984. Soc. Detecting insect larvae in fruit by vibrations produced. The fruit flies of the genus Anastrepha. Entomol. J. 177. 169. De­ scription of immature stages of An­ astrepha interrupta. Entomol. Landolt PJ. 60: 1 116 1 8 7 . J. Sci. 1977. 196 1 . 71 pp.) in Costa Rica. Wharton RA. 1 976. J. Entomol. by University of Guelph on 10/10/12. Entomol. Descrip­ (Mangifera indica L. serpentina (Diptera: Teph­ ritidae). 1 942. Entomol. Agric. Steck GI. 1984. Airplane applications of malathion bait spray for Mexican fruit fly control. Publ. 439: 1-1 1 2 189. Lopez PD. . Brasil. A review of research done with the Mexican fruit fly and the citrus blackfly in Mexico by the entomology research division. Entomol. Am. Am. Soc. Trujillo PG. 1987. cultivar Francis. Pest Manage. Benner Je. 1 946. J. 2:3-13 Sivinski J. 79:75-8 1 186. 166. The analysis and identification of sounds produced by the male Caribbean fruit fly. 1 162-67 176. Turrialba 37: 245-5 1 180. Environ. Am. Epsky N. 174. 1990. Health A19 3:367-75 190. Soc. J. 19: 1491-95 Sivinski J. 1 99 1 . Warthen ID. 1988. Spishakoff LM. 1990. Sanchez-Riviello M. J. and key to 13 species. 1967. Entomol. Methods for identification of An­ astrepha larvae (Diptera: Tephritidae). 35 pp. 1970. 178. Sivinski J. Two new tropical fruit flies of the genus Anastrepha. 1990. Econ. McDow 11. Ann. Fkiooen-Andersen JL. Car­ yologia 43:229-41 Solferini VN.oes entre bacterias e Anastrepha (Diptera: Tephritidae). 8 1 : 1004-9 183. 167. Am. 172. Am. Webb Je.39:155-178. 168. Proc. 196 1 . Sanchez-Riviello M. Development of im­ mature stages of Anastrepha serpentina in relation to temperature. Genet. Webb JC. Insect Behav. En­ tornol. 69:41 5-20 1 9 1 . 1988. Stud­ ies on the population dynamics of the fruit flies. Econ. Interar. Brasil. 1990. L6pez PD . Morgante JS. Litzkow C. J. Entomol. Wash. 60:992-94 Sivinski J. Burk T. Soc. Calkins CO. Colored spherical traps for capture of Caribbean fruit fly. limae. 73: 123-28 Sivinski J. 1994. Soto-Manatiu J. Solferini VN. Anastrepha suspensa (Loew). Guillen-Aguilar J. Entomol. liron LF. USDA Misc. Dispersal and migration of tepa-sterilized Mexican fruit flies. Heath RR. Biochemical system­ atics and population genetic structure of Anastrepha fraterculus and related species (Diptera: Tephritidae). Morgante JS. 1989. Agric.ANASTREPHA BIONOMICS AND MANAGEMENT Tephritidae) larvae in mango. 188. PhD thesis. Pheromone deposition on leaf-territo­ ries by male Caribbean fruit flies (An­ astrepha suspensa) (Loew). 16:186-93 Shaw JG. Insect Behav. 173. 8 1 :406-9 Shaw JG. Steck GJ. Soc. Entomol. Entomol. For personal use only. and A . Annu. Chambers DL. 1993. Econ. Acoustic courtship signals in the Car­ ibfly Anastrepha suspensa. Rev. Chemical control and eco­ logical observations of fruit flies of the genus Anastrepha Schiner (Diptera: Tephritidae) on mango. X\X\XiX2:X\X2Y mechanism of sex determination in Anastrepha bistrigata and A .annualreviews. 35:425-27 170. Hernandez R. tion of immature stages of Anastrepha bistrigata (Diptera: Tephritidae) . Res. 8 1 :994-1003 184. Anastrepha 175. 179. Exploratory studies with soil toxicants to control the Mexican fruit fly. 185. Celedonio­ Hurtado H. Acoustical behavior and sexual success in the Caribbean fruit fly. 1984. with notes on generic synonymy (Dip­ tera: Tephritidae). Carroll LE. Uebel EC. J. Karyotype study of eight species of Anastrepha (Diptera: Tephritidae). Webb JC. Stokes J B . In press Solferini VN. 72:265-76 Shaw JG. Steyskal GC. Soc. 54:600-1 Shaw JG. Univ. 171. Entomol. Fla. 92:333-46 182. Rev. Trop. Sharp JL. Proc. Steyskal GC 1977. Steyskal GC. Malavasi A. Lekking and the small-scale distribution of the sexes in the Caribbean fruit fly Anastrepha sus­ pensa (Loew) . Anim. Jacobson M. Sex­ ually dimorphic developmental rates in the Caribbean fruit fly (Diptera: Tephritidae). Downloaded from www. Sao Paulo. 32: 101 1-16 Sivinsky J . 3 1 : Stone A . Spec. Wash. 84: 10-28 1 8 1 . Behav. History and use of the McPhail trap. Ann. A. Jiron LF. Fla.

13:650-56 192. Entomol. Tephritidae) assinaladas no Brasil. Ramadan MM. 1979. Entomol. Brasil. (Diptera: Tephritidae) in Costa Rica. Univ. 1992. Classical biological control of fruit-infesting tephritidae. 23: 35-41 198. Hart WG. Rhode RH. Brasil. 1:2-7 196. Mcinnis DO. Sao Paulo. 1983. 105 pp. Environ. 19868 (Diptera: Tephritidae). 610 pp. 154a.39:155-178. Augmentative releases of Annu. 1989. See by University of Guelph on 10/10/12. UK: CAB International. Hawaii. Downloaded from www. White 1M. Oxon. 16-30 Diachasmimorpha tryoni (Hymenop­ . Zucchi RA. suspensa (Loew) (Diptera: Teph­ ritidae). Bioi. pp. 1968 (Diptera. 3B:303-13 193. See Ref. 197. Herr tera: Braconidae) to suppress a Medi­ terranean fruit fly (Diptera: Tephritidae) population in Kula. 1978.annualreviews. Entomophaga 26:285-90 194. Zwolfer H. 42a. Fruit Flies o f Economic Importance: Their Identification and Bionomics. 1981. Entomol. 195. Contr. Wong TTY.178 ALUJA JC. Fischel MM. Gilstrap FE. Elson-Harris MM . Rev. Wharton RA. Rev. Taxonomia das es­ pecies de Anastrepha Schiner. Nishimoto JI. Life systems and strategies of resource exploitation in tephritids. For personal use only. Hyme­ nopterous egg-pupal and larval-pupal parasitoids of Ceratitis capitata and Anastrepha spp. Mochizuki N. Zucchi RA. PhD thesis. 1994. Novas especies de Anastrepha Schiner . Wharton RA . 1991. Maui.