Natural Enemies of True Fruit Flies (Tephritidae) - USDA - aphis

United States 

Department of 

Agriculture 

Marketing and 

Regulatory 

Programs 

Animal and 

Plant Health 

Inspection 

Service 

Plant Protection 

and Quarantine 

Natural Enemies of 

True Fruit Flies 

(Tephritidae) 

Jeffrey N. L. Stibick 

Psyttalia fletcheri (shown) is the only fruit fly parasitoid introduced into Hawaii 

capable of parasitizing the melon fly (Bactrocera cucurbitae) 

United States Department of Agriculture 

Animal and Plant Health Inspection Service 

Plant Protection and Quarantine 

4700 River Road 

Riverdale, MD 20737 

February, 2004


Introduction 

Introduction 

Fruit flies in the family Tephritidae are high profile insects among 

commercial fruit and vegetable growers, marketing exporters, 

government regulatory agencies, and the scientific community. 

Locally, producers face huge losses without some management 

scheme to control fruit fly populations. At the national and 

international level, plant protection agencies strictly regulate the 

movement of potentially infested products. Consumers throughout 

the world demand high quality, blemish-free produce. Partly to satisfy 

these demands, the costs to local, state and national governments are 

quite high and increasing as world trade, and thus risk, increases. 

Thus, fruit flies impose a considerable resource tax on participants at 

every level, from producer to shipper to the importing state and, 

ultimately, to the consumer. (McPheron & Steck, 1996) 

Indeed, in the United States alone, the running costs per year to 

APHIS, Plant Protection and Quarantine (PPQ), (the federal Agency 

responsible) for maintenance of trapping systems, laboratories, and 

identification are in excess of US$27 million per year and increasing. 

This figure only accounts for a fraction of total costs throughout the 

country, as State, County and local governments put in their share as 

well as the local industry affected. If a emergency program is mounted 

against some exotic fruit fly invasion, the costs to PPQ increase again, 

and may be as much as an additional $20 million per year. An 

exceptional program, such as the Medfly eradication Program in 

California in 1980-82 required the expenditures of $100 million for 

this one program alone, over roughly 3 years, for all parties involved 

(Schribner, 1983). Costs in other areas of the world are no less. For 

Japan, the cost of eradication of the Oriental Fruit Fly from its 

south-western islands in the 1980’s was US$32 million (White and 

Elson-Harris, 1992). 

It goes without saying that the discovery of exotic fruit flies in a 

country result in labor intensive and very costly eradication programs 

from time to time, whenever a pest is discovered. This is especially 

true with the growth and globalization of world trade, when fruit 

produce and their associated pests are carried around the world at 

speeds unheard of before the nineteenth century. 

During the development of a New Pest Response Guidelines for The 

Economic Fruit Flies, Family Tephritidae, by USDA, APHIS, PPQ, a 

comprehensive list of natural enemies of Tephritids was developed. 

This was recognized as a valuable source of information for those 

interested in classical or augmented biological control for fruit flies in 

the future. Since the work is still in preparation, the information is 

considered apart from other means of control that are discussed in the 

parent document. 

3-2 Natural Enemies of True Fruit Flies 02/2004-01 

PPQ


Classical Biological Control 

This paper provides, as far as is possible, the natural enemies of fruit 

flies throughout the World. It is limited to fruit pest species, as 

opposed to leaf, stem and root pest species, following White and 

Elson-Harris, 1992. 

Classical Biological Control 

A basic premise of classical biological control is that the target pest 

will be brought under some degree of control while nontarget 

organisms are not affected. To do this, the introduced bio-control 

agent must be relatively host specific. 

Often, this can only be determined after release of the agent into the 

environment.To remedy this dilemma, lab studies attempt to 

determine the physiological host range of the agent in order to predict 

the ecological host range as well. This does not always work when 

nontarget possible hosts are exposed to the agent under the confined 

circumstances of the lab. Many conditions in the outside environment 

determine which possible hosts are attacked, such as spatial or 

temporal overlap, host ranges and/or substrates of target and 

nontarget species, temperature and humidity, tolerances, and others. 

(Solter, et al, 1997; Hajek, et al, 1996) 

To the extent possible, biocontrol agents should be carefully 

considered for their possible impact on nontarget organisms. Some 

general rules are as follows: 

A. Predators are usually more generalist than parasites. 

B. The known host specificity of an agent, including information 

on behavior of related taxa. 

C. Selection and release of agents known to be host specific or 

attack only closely related non-target species. 

To this end, information on the available Biological Control organisms 

are given in table form to allow comparisons between different 

Tephritid species as a decision-making tool, and to help in the 

selection of the best combination of useful biological control agents. 

Microorganisms/microbial toxins - TABLE 1 

Table 1 lists the micoorganisms/micobial toxins used against the fruit 

flies species listed therein. Biological Modes are given separately 

under each species. The Products or Microbiol Pesticides 

(Biopesticides), are consecutively numbered under each Biological 

02/2004-01 Natural Enemies of True Fruit Flies 3-3 

PPQ


Microorganisms/Microbial Toxins Used Against the Tephritidae - Table 1 

Mode. The use of such techniques are generally considered as 

Biotactical techniques, but is included in this document as the 

products may be considered as coming from natural enemies. 

It should be remembered that nongenetic resistence may take place. 

This includes phenotypic changes in insect behavior or physiology and 

of host plant interference with pesticide action, including microbial 

pesticides such as entomopathic bacteria and viruses. These are 

particularly sensitive to plant chemistry because they infect through 

the gut. As a consequence, the composition of foliage ingested with 

the microbial pesticide can dramatically influence its effectiveness. 

(Appel & Schultz, 1994) 

Another factor to consider is rainfall. It has been suggested that a 

light rainfall may help in prolonging the period of activity of viral 

preparations by moving the virus downwards, towards the more 

shaded parts of a plant and away from light. This would help to 

prolong its effectiveness. No absolute proof of this hypothesis has yet 

been made. (D’Amico & Elkinton, 1995) 


Species Biological Mode Product Specif ics 

Anastrepha 

fraterculus 

Anastrepha 

ludens 

Through 

transovarian 

transmission 

Through adult 

consumption 

Immersion in a 

conidial 

suspension 

An Endosymbiotic bacteria of the genus 

Wolbachia. 

Agent:Phloxine B-Mazoferm 802 

formulation for field application 

Mazoferm 802 - corn condensate 

hydrolyzed by a Lactobacillus sp. 

Phloxine B - photoactive dye used in 

formulations of foods or 

pharmaceuticals and cosmetics 

Laboratory strain of Anastrepha sp. 2 

aff. fraterculus. from Brazil . No 

evidence of alterations in the 

reproductive ability of A. fraterculus 

(Selivon, et al., 2002) 

In field trials, aerial application 

reduced the population of mexfly by 

85% against the control catch 

(Moreno, et al., 2001) 

A fungus, Metarhizium anisopliae. Laboratory test, which resulted in 

mortality rates from 37.9 to 98.75%. 

Also a conidial suspension (2.5 X 

106 UFC.ml) was applied to a soil 

surface, resulting in reduction in 

adult emergence of 22-43% 

(Lezama-Gutierrez, et al., 2000) 


PPQ

Anastrepha 

obliqua 

Bacillus 

thuringiensis, a 

gram-positive, 

spore-forming soil 

bacterium fed to 

larvae & adults. 

Paratransgenesis 

in which genes 

and proteins of 

selected species 

of the microbial 

fauna of Mexfly 

guts are altered, 

and fed to adults. 

Beauveria 

bassiana, a 

fungus, fed to 

larvae and adults. 




Anastrepha 

obliqua (Cont.) 

Anastrepha 

serpentina 

Anastrepha 

suspensa 

Agent: beta exotoxin toxic towards 3rd 

instar larvae 

Exposed in lab only (Toledo, et al, 

1999) 

Stablility of pZeoEGFP, a plasmid. Stability suggests that derivatives of 

this plasmid could serve to introduce 

heterologous genes and proteins, 

that, when ingested by adults, such 

proteins/products would be delivered 

directly to the gut of host flies. If 

toxic, this would serve to control fly 

populations. If nutrition is enhanced, 

they could improve vigor of colony 

flies for SIT (Peloquin, et al., 2000; 

Kuzina, et al., 2001). 

An entomopathogenic soil-borne fungus 

that attacks many different insects and 

exists in many different strains. 

Airborne conidia An unidentified fungus in the 

Entomophthorales 

Through adult 

consumption 

Bacillus 






Bacillus 






Plant growth 

regulator 

Agent:Phloxine B-Mazoferm 802 








instar larvae 


instar larvae 

A hormone from the fungus Gibberella 

fujikuroi, gibberellic acid (GA 3) + a 

surfactant (L-77) Commercial 

formulations available 

In Laboratory, most toxic to adults @ 

rates ranging from 98 to 100%. 

Larvae were affected @ rates ranging 

from 2 to 8%. Doses ranged from 

1.0 x 10 8 to 1.6 x 10 8 (Rosa, 2002). 

Capable of wiping out laboratory 

populations via airborne conidia 

(Baker, et al., 1944). 


reduced the population of medfly by 




1999) 


1999) 

Reduces host susceptibility/ 

attractiveness to fruit flies by 

preventing aging of the peel 

(yellowing) 

10 ppm GA3 /L-77 sprayed to run-off 

on fruit (Greany et al, 1991) 


PPQ




Anastrepha 

spp. 

Bactrocera 

ascita sp. B 

Bactrocera 

cucurbitae 

Bactrocera 

cucurbitae 

(Cont.) 

Bactrocera 

carambolae (= 

B. dorsalis sp. 

A) 

Bactrocera 

caudata 

Bactrocera 

diversa 

Through larval 

contact 

Saccharopolyspora 

spinosa, a soil 

bacterium, through 

ingestion. 

Through 

transovarian 

transmission 

Through adult 

contact (?) 

Through adult 

contact 

Through adult 

contact 


contact 

Through adult 

contact 

Through adult 

contact 

Through adult 

contact 

Through adult 

contact 

An Entomogenous nematode Agent: 

Steinernema carpocapsae 

Spinosad 

Agent: Spinosyn A Spinosyn D 

+ 

SolBait 


Wolbachia. 

Augmentation to eradication is 500 

nematode per cm 2 applied to soils. 

Results similar to fruit flies in Hawaii 

(Lindegren, 1990) 

Test plots sprayed through aerial 

applications at the rate of 1.8 liters/ 

hectare resulted in 54% reduction in 

Caribfly populations. At 3.5 liters/ 

hectare the reduction was 73%. 

(Burns, et al., 2001) 

No evidence of alterations in the 

reproductive ability of A. suspensa 

(Werren, et al., 1995) 

unspecified fungi in the Laboulbeniales Affect various body parts, but 

apparently little inconvenience to the 

host (Baker, et al., 1944) 


Wolbachia, inducing cytoplasmic 

incompatibility, thelytokous 

parthenogenesis, male-killing or 

feminization. 

Agent: Avermectin B1 (MK-936) 

a powerful toxin derived from 

Streptomyces avermitilis fermentation 

(EPA, 1989) 







feminization. 





feminization. 





feminization. 





feminization. 

In field collected adults from 

Thailand (Jamnongluk, et al, 2002) 

In Lab, applied to thorax, at 0.042 

ug/g LD 50 15 days 

With sublethal doses, fecundity, and 

fertility reduced by up to 79% 

(Albrecht & Sherman, 1987) 



Causes 89% mortality (Lindegren, 

1990) 










PPQ

Bactrocera 

dorsalis 

Bactrocera 

dorsalis (cont.) 

Bactrocera 

dorsalis sp. A1 

Opius oophilus, a 

Parasite 

Through adult 

contact. 

Through rotting 

fruit by contact 

with larvae and 

pupae 


contact. 

Through adult 

consumption 

Through adult 

contact 




Bactrocera 

modica 

Bactrocera 

oleae 

Through adult 

contact 

Lasioptera 

berlesiana, a 

Dipterous 

predator. 


siniosa, a soil 

bacterium.Through 

larval contact. 

Cricket virus, feed 

to the adult, 

infection of gut 

and cross-infection 

from one fly to 

another by 

fecal-oral route 

(fecal 

contamination of 

food and liquid 

sources). 

unspecified bacteria and fungi 

(EPA, 1989) 

Agent: Leucothix mucor 

Agent: Avermectin B 1 (MK-936) 

a powerful toxin derived from 

Streptomyces avermitilis fermentation. 

Females serve as carriers of bacteria 

and fungi that destroy a considerable 

portion of the host eggs (Clausen, 

1956) 

In Lab, applied to thorax, at 0.021 

ug/g LD50 15 days 

With sublethal doses, fecundity, ard 

fertility unchanged (Albrecht & 

Sherman, 1987) 

A Bacterial pathogen Identified as a major cause of 

mortality. (Gingrich, 1993) 

An Ent Agent: Steinernema carpocapsae 

an entomogenous nematode 


nematode per cm2 applied to soils. 


1990) 

In lab trials, males were killed in 

less than 2 hours (Liquido, et al., 

1995) 

Agent: 3.6% Phloxine B 

+ 

7.1% methyl eugenol 

An Endosymbiotic bacteria of the genus In field collected adults from 





feminization. 

An Endosymbiotic bacteria of the genus In field collected adults from 





feminization. 

Macrophoma dalmatica, a fungus Unknown 

(Narayanan & Chawla, 1962) 

Spinosad 

Agent: Spinosyn A 

Spinosyn B 

A picornavirus from Crickets 

Agent: CrPV 

Exemption for use in a proteinaceous 

bait spray 

(EPA, 1999) 

Highly pathogenic to D. olea: 50% 

within 5 days,80% within 16 days. 

(Gingrich, 1993; Manousis & Moore, 

1987) 


PPQ




Bactrocera 

pyrifoliae 

Bactrocera 

tryoni 

Ceratitis 

capitata 

Infects pupae in 

laboratory and 

natural 

populations. 

Bacillus 






Through adult 

contact 

Queensland fruit 

fly virus (QFFV), 

through adult 

contact. (Moussa, 

1978) 

Through adult 

contact 

Through adult 

consumption 



bacterium.Through 

larval contact. 

A Picorna-like virus Infestation in several stocks of 

laboratory and natural populations in 

Greece (Knowles, 1998; Manousis, 

et al., 1987) 

Isolates TE 37.18 and Mex 63 gave a 

mortality of over 50% in adults. 

Isolate JC 120 gave a mortality of over 

69.2 to 95% in larvae. 

Strain A4 with 140kDa and 32kDa have 

insecticidal activity for olive fly. 





feminization. 

A picornavirus, probably the 

Picornaviridae, in the genus Enterovirus. 

Agent: QFFV, easily propagated through 

infestations of laboratory populations 

(Bashiruddin, et al., 1988) 

Agent: Avermectin B 1 (MK-936), a 

powerful toxin derived from 

Streptomyces avermitilis fermentation 

(EPA, 1989) 

Agent:Phloxine B - Mazoferm 802 







Agent Phloxine B - Mazoferm 802 

+ 

Fopius arisanus (a parasite population in 

test area 

Spinosad 


Spinosyn B 

+ 

Fopius arisanus (a parasite population in 

test area 

Infestation in both labortory and 

natural stocks in Spain. (Alberola, 

et al., 1999) 

Labortory testing only (Sivropoulou, 

et al., 2000) 



Infected, probably through feeding, in 

the midgut.Highly pathogenic, 

resulting in a mortality of 40-50% in 

2nd and 3rd week after emergence 

of adults.Surviving adults may have 

reduced fecundity for a few days 

(Bashiruddin, et al., 1988) 

In Lab, applied to thorax, at 0.29 ug/ 

g LD50 10 days 

With sublethal doses, fecundity, ard 

fertility reduced by up to 29% 

(Albrecht & Sherman, 1987) 


reduced the population of medfly by 



Weekly ground spraying 

Measured populations only (Vargus, 

et al., 2001) 

Weekly ground spraying 

Measured populations only (Vargus, 

et al., 2001) 


PPQ

Ceratitis 

capitata 

(Cont.) 


spinosa, a soil 

bacterium, through 

ingestion. 

Entomophthora 

muscae, an 

entomogenous 

fungus 

Entomophthora 

schizophorae 

Through 

adsorbtion into the 

peel of the fruit 

Through larval/ 

pupal contact. 





prepupal contact. 


prepupal/pupal 

contact. 


contact. 

Through contact 

with the adult. 


with the pupa 

Through egg 

contact 


with the pupa 

Spinosad 


Spinosyn D 

+ 

SolBait 

Kills the adult through growth and 

sporulation. 

Kills the adult through growth and 

sporulation. 

A hormone from the fungus Gibberella 

fujikuroi, gibberellic acid (GA 3 ) + a 

surfactant (L-77) 

Commercial formulations available 

An Entomogenous nematode 

Agent: Steinernema feltiae 




Steinernema riobrave 

A predator nemetode 

Agent:Diplogaster sp. 

An exotoxin product Agent: Bacillus 

thuringiensis 

A fungal Agent: Metarhizium anisopliae 

var. anisopliae 

A bacterial Agent: Serratoa marcescems 

strain (EC 3.1.21to EC 3.1.31) 

A microsporidian Agent: Nosema 

tephritidae 

Test plots sprayed through aerial 

applications at the rate of 1.8 liters/ 

hectare resulted in 96% reduction in 

Medfly populations. At 3.5 liters/ 

hectare the reduction was 99%. 

(Burns, et al., 2001) 

Field collection of adults only (Uziel & 

Yuval, 2003) 

Field collection of adults only (Uziel % 

Yuval, 2003) 

Reduces host susceptibility/ 

attractiveness to fruit flies by 

preventing aging of the peel 

(yellowing) 

10 ppm GA3/L-77 sprayed to run-off 

on fruit (Rossler & Greany, 1990; 

Greany et al, 1991) 

Optimal 500 nematode per cm2 as a 

soil drench. Causes 87.1% 

mortality. (Lindegren, et al, 1990) 




1990) 

Optimal 100 nematode per cm 2 

applied to soils. Causes 82% 

mortality (Gazit, et al., 2000) 

Test shows nematodes will devour 

larva by paralysis and digestion of 

contents. (Osman, et al.,1990) 

Also lethal to mammals if injected 

and not registered for use in many 

countries, including the U.S. 

(Gingrich, 1993) 

At applications of 8.25, 7.8 and 28 x 

10 6 conidia/ml, deaths were about 

6-33.7% in the first week to 

24.1-66.3% in the second week and 

100% by the 6th week (Garcia, et al., 

1989). Through soil inoculation 

(Ekesi, et al., 2002) 

Prevented eggs from hatching under 

lab conditions (Moore & Nadel, 

1961) Dosage? 

Virulent (Steiner & Mitchell, 1966) 

Dosage? 


PPQ




Ceratitis 

capitata 

(Cont.) 

Ceratitis 

cosyra 

Ceratitis rosa 

fasciventris 

Dacus 

destillatoria 

Rhagoletis 

cerasi 

Rhagoletis 

cingulata 


with adult 

A bacterial Agent: Beauveria bassiana A facultative pathogen attacking 

adults. (Mitchell & Saul, 1990). 

Through soil inoculation (Ekesi, et 

al., 2002) 

In the adult A Picornavirus Agent: Ceratitis 

Picornavirus V 

In lab, a natural virus found in the 

species. Pathogeny questionable 

(Plus &Cavalloro, 1983; Bashiruddin, 

et al., 1998; Knowles, 1998) 

In the adult A Reovirus Agent: Ceratitis Reovirus I In lab, a natural virus found in the 

species. Pathogeny questionable 

(Plus & Cavalloro, 1983) 

In the adult A Rhabdovirus Agent: Rhabdovirus signa In lab, pathogenic when injected into 

adult. (Plus & Cavalloro, 1983) 

In the adult A Picornavirus Agent: Picornavirus C 

(DCV) 


with 3rd instar 

larvae and pupae 


with the pupa 


with 3rd instar 

larvae and pupae 


with the pupa 

Through adult 

contact 

Maternally 

Inherited 



bacterium. 

In lab, pathogenic when injected into 

adult. (Plus & Cavalloro, 1983) 


larvae, pupae and deferred mortality 

of emerging adults through soil 

inoculation (Ekesi, et al., 2002) 



A fungal agent attacking larvae, 

pupae and deferred mortality of 

emerging aduits through soil 



larvae, pupae and deferred mortality 

of emerging adults through soil 








feminization. 



incompatibility. 

Success Agent:Spinosyn A 

Spinosyn B 

A fungal agent attacking larvae, 

pupae and deferred mortality of 

emerging aduits through soil 




Europe. Suggested introduction of 

cytoplasmically incompatibile males 

to an invading population of R. cerasi 

(Riegler, 2002). 

Treat at .062-.125 lb/acre ai at a 

rate of 4-8 fl oz/acre or 1-2 fl oz/100 

gal of dilute spray. For Stone Fruit in 

California (Dow AgroSciences, 1999) 


PPQ


Rhagoletis 

completa 

Rhagoletis 

indifferens 






fauna of fruit fly 








fauna of fruit fly 





bacterium. 

Stablility of pZeoEGFP, a plasmid. 

Enterobacter agglomerans, a symbiotic 

bacterium in the gut, through adult 

consumption. 

Stablility of pZeoEGFP, a plasmid. 

Kiebsiella pneumoniae, a symbiotic 

bacterium in the gut, through adult 

consumption. 

Success Agent: 

Spinosyn A 

Spinosyn B 


Parasites and Predators - TABLE 2 

Stability suggests that derivatives of 









flies for SIT (Peloquin, et al., 2000) 

Stability suggests that derivatives of 









flies for SIT (Peloquin, et al., 2000) 

Treat at 0.062-.125 lb/acre ai at a 

rate of 4-8 fl oz/acre or 1-2 fl oz/100 

gal of dilute spray. For Stone Fruit in 

California (Dow Agro Sciences, 1999) 


Biological control of fruit flies has not always been regarded as 

successful when measured by economic returns, partly because the 

economic threshold for fly damage is very low. In addition, a lack of 

information on the nontarget impact of parasites has drawn concern 

from environmentalists and conservation biologists (Duan & Messing, 

1977). For these reasons, classical biological control for fruit flies 

seemingly suffers from a commonly encountered set of biological 

difficulties; especially: 

1. low fecundity of parasitoids compared to fruit flies, 

2. poor tracking of fly populations by parasitoids, due either to 

relatively ineffectual foraging at low host densities or poor 

survival through periods when fruit and hosts are absent, and 

3. Host refuge from parasitoid attack by fly larvae in large/ 

thick-skinned fruit or under dense litter for pupation. 

(Sivinski, 1996) 

4. Some species of parasites may exhibit diapause. If they are used 

for parasitoid mass rearing and release programs such species 

could complicate handling procedures and raise costs (Aluja, et 

al., 1998) 


PPQ



5. Impacts of introduced parasites on nontarget organisms (Duan & 

Messing, 1977) 

The solution to the first two problems is comparatively simple: 

augment the numbers of parasitoids at critical times and places and 

use the technique in conjunction with other controls, notably sterile 

release. (Sivinski, 1996) 

The third problem may be partly addressed by employing parasites, 

especially late instar or pupal parasites, which forage extensively over 

fallen/broken fruit or even enters damaged fruit crack and crawls 

through the pulp. Again, such parasites should be augmented. 

(Sivinski, 1996) 

The fourth problem may be addressed by employing non-diapausing 

parasites or to avoid exposing parasites to stimuli eliciting diapause or 

selecting lines that reduce incidence of diapause. (Aluja, et al., 1998) 

The fifth problem has been shown to be partly non-existant by Duan & 

Messing (1977, 2000), who demonstrated that in Hawaii, at least, such 

parasites had no harmful effect on flowerhead-infesting fruit flies and 

probably limited effect on gall-forming and other groups of tephritids. 

The usefulness of certain particularly effective predators should also 

not be overlooked, especially if one can be obtained and released in 

large numbers. 

In addition, certain newer techniques, such as augmentative releases 

of parasites, selected specific predators, and biopesticides, such as 

those listed in Table 1, may well bring the pest population down to 

noneconomic levels if carefully planned. 

Table 2 lists the known parasites and predators of the fruit pest 

species of the Tephritidae. They are given under the Tephritid species 

involved, with such notes from the literature that are available. While 

an effort has been made to keep up with the literature regarding 

synonymous names, there is no guarantee that these names will still 

apply to the parasite or predator in question. 

Limitations to this list include restrictions on the number of citations 

given for each parasite or predator, since a repeat of citations serve no 

useful purpose. Generally, a citation would have to add some useful 

information or fact about a given parasite or predator. Also, if 

parasitoid guilds were mentioned without the fruit fly host, this would 

lead to a painstaking task to determine the correct species/host 

relationship and this task has been avoided. Other, more specific 

references may fill in this gap. 


PPQ



In addition, due to time constraints, there is no guarantee that all 

parasites of the listed economic species are given here, although a 

reasonable effort has been made. Parasites of many noneconomic 

fruit flies were included in order to provide reasonable coverage of 

most of the known Tephritid parasites as an aid in program efforts. 

The cross-introduction of many parasites from one area to another 

and their successes and failures in over a century of such attempts is 

barely mentioned, and only when such introductions were successful. 

One of the factors in successful or unsuccessful use of parasites deals 

with the fact that the rate of parasitization may vary with the types of 

hosts infested by fruit flies. That is, the attack response of the 

parasitoids may be highly discriminating among the different hosts of 

a particular fruit fly pest. For Biosteres arisanus, for example, certain 

fruit like banana or coffee are more preferred by this parasitoid than 

citrus for attack on fruit flies (Bautista & Harris, 1996). 

Under NOTES, a very brief line or two may provide some information 

about the parasite or predator which may be useful if the species is to 

be considered for program purposes. In all cases, the references are 

cited in this column. 

Some economic species are listed without any parasites or predators. 

This serves the purpose of highlighting that these species need further 

foreign exploration. No known noneconomic species are so listed. 

A good background document on the use of parasites for control of 

fruit flies is “A Review of Mediterranean Fruit Fly Parasites with special 

Reference to Western Australia” (Hardie, 1982). This paper is much 

more than the title implies, as there is a lot of useful information and 

references to aid in any program dealing with release of parasites. 

Wharton, 1989, gave a good summary of classical biological control 

efforts, pointed out some of the reasons for the many failures and gave 

suggestions for improving future efforts.These items are summarized 

below: 

Early Failures of classical biological control of fruit flies 

1. Transportation difficulties 

2. Rearing difficulties 

3. Emphasis on laboratory colonization 

4. Exceptionally small numbers 

5. Dropping species as a result of unsuccessful attempts 

6. Study of the biology of natural enemies lacking 


PPQ



7. Failure to adequately document efforts, no prerelease surveys, 

lack of published records, and failure to properly evaluate 

releases 

Improving future classical biological control efforts 

1. Retesting, with improved rearing methods, those species which 

were unsuccessful in the past 

2. Trying, with improved rearing methods to raise more difficult 

parasitoids that were avoided in favor of easily reared species 

3. Make more direct releases for field evaluation, rather than 

emphasis on laboratory colonization 

4. Complete studies on the biology of the natural enemies to be 

utilized 

5. Collect data to document impact of these parasitoid releases in 

reducing the targeted fruit fly population densities 

There has been a gradual shift from classical biological control to 

augmentative biological control with mass rearing and the inundative 

release approach (Wong, et al., 1991; Gingrich, 1993). This effort may 

address some of the perceived shortcomings of classical biological 

control in the context of a program effort for dealing with an invasive 

pest and it does seem to have its advantages in dealing with a massive 

pest population that is thriving in a new environment, by causing that 

population to diminish rapidly or even crash. The purpose of releasing 

such numbers is: 

A. To eradicate the invading population. 

B. To suppress the invading population over time and be self 

sustaining 

C. To control the invading population in as low numbers as is 

possible and to augment biological control with sterile 

releases of the fruit fly and/or releases of micoorganisms. 

For this purpose, Ashley and Chambers (1979) did a study on 

maximum rearing efficiency of Biosteres longicaudatus, on early 

Anastrepha suspensa 3rd instar larvae. They concluded that with 

ovipositional cages of size 24.5 cm 3 , maximum efficiency is reached at 

a parasite density of 125 pairs of this species per cage, producing 

1,896 progeny per 24 hr. exposure period. 

Other approaches such as the release of pupal parasites to attack any 

pupae of the target pest that survive other program efforts, such as 

sterile release, biopesticides or other biotactical measures and/or 

augmented parasitic releases of egg and/or larval parasites or 

multiparasitism have been discussed. (Wong, et al., 1992; Bautista & 


PPQ



Harris, 1997; Sivinski, et al., 1998). Augmented release follows this 

table. Biotactical control is presented in The New Pest Response 

Guidelines for the Economic Fruit Flies mentioned previously. 


PPQ


Parasites and Predators of the Tephritidae - Table 2 


Species Parasites/Predators Notes 

Acidiella 

angustifascia 

Bracon intercessor A Braconid parasite from Russia, China, Korea (Kandybina, 1977). 

Opius Myolejae A Braconid parasite from Russia, China, Korea (Kandybina, 1977). 

Opius Mariae A Braconid parasite from Russia, China, Korea (Kandybina, 1977). 

Acanthiophilus Opius afreutretae A Braconid larval-pupal parasite from South Africa (Narayanan & Chawla 

muiri 

(1962). 

Acidiella 

angustifascia 

Bracon intercessor A Braconid parasite from Russia, China, Korea (Kandybina, 1977). 

Opius myolejae (?) A Braconid parasite from Russia, China, Korea (Kandybina, 1977). 

Opius mariae A Braconid parasite from Russia, China, Korea (Kandybina, 1977). 

Acidiella 

echinopanacis 

Opius mariae A Braconid parasite from Russia, China, Korea (Kandybina, 1977). 

Anastrepha spp. Biosteres 

A Braconid larval-pupal parasite introduced into Costa Rica (Wharton, et al., 

longicaudatus 1981); into Mexico Peru, and Bolivia (Rogg & Camacho, 2003). 

Biosteres oophilus A Braconid egg-pupal parasite introduced into Costa Rica (Wharton, et al., 

1981) 

Dicerataspis flavipes An Eucoild larval parasite from Brasil (Guimaraes, et al., 1999). 

Doryctobracon A Braconid larval-pupal parasite from Costa Rica, Eucador, Venezuela 

areolatus 

(Wharton, et al., 1981; Molineros, et al., 1992; Katiyar, et al., 1995). 

Doryctobracon cereus A Braconid parasite from Peru and Bolivia 

Doryctobracon A Braconid larval-pupal parasite from Costa Rica to Ecuador; widespread, 

crawfordi 

high parasitism (36+%) (Wharton, et al., 1981; Molineros, et al., 1992). 

Doryctobracon zeteki A Braconid larval-pupal parasite from Costa Rica (Wharton, et al., 1981). 

Opius anastrephae A Braconid larval-pupal parasite from Costa Rica (Wharton, et al., 1981). 

Opius bellus A Braconid larval-pupal parasite from Costa Rica (Wharton, et al., 1981). 

Ganaspis carvalhoi A Cynipid larval-pupal parasite from Costa Rica (Wharton, et al., 1981). 

Odontosema 

A Cynipid larval-pupal parasite from Costa Rica, Puerto Rico (Wharton, et al., 

anastrephae 

1981; Thompson, 1943). From Brasil (Guimaraes, et al., 1999). 

Aceratoneuromyia 

indica 

A Eulophid larval-pupal parasite from Costa Rica (Wharton, et al., 1981) 

Ganaspis sp. A Cynipid parasite from Puerto Rico. 

Acari Unspecified mites from Mexico (Baker, et al., 1944). 

Asobara anastrephae A solitary Alysiid entoparasitoid from Brazil (Leonel, et al., 1995). 

Opius sp. A Braconid solitary entoparasitoid from Brazil (Leonel, et al., 1995). 

Pachycrepoideus 

dubius 

A Pteromalid parasite (Narayanan & Chawla, 1962). 

Trichopria 

anastrephae 

A Diapriid parasite from Brazil (Narayanan & Chawla, 1962). 

Aganapsis pelleranoi A Eucoilid parasite from Venezuela (Katiyar, et al., 1995). From Brasil 

(Guimaraes et al., 1999). 

Lopheucoila 

A larval-pupal Eucoilid parasite from Mexico, Central and South America 

anastrepha 

(Gallardo & Diaz, 1999) 


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Anastrepha 

alveata 

Anastrepha 

amita 

Anastrepha 

antunesi 

Anastrepha 

bahiensis 

Anastrepha 

benjamini 

Anastrepha 

bistrigata 

Doryctabracon 

areolatus 



A Braconid larval-pupal parasite of late instars from Mexico to Argentina 

(Sivinski, et al., 1997; Piedra, et al., 1993; Lopez, et al., 1999; Sivinski, et 

al., 2000; Aluja, et al., 2000). 

Utetes anastrephae A Braconid larval-pupal parasite from Mexico (Lopez, et al., 1999; Sivinski, et 

al., 2000). 

Opius hirtus A Braconid larval-pupal parasite from Mexico (Lopez, et al., 1999; Sivinski, et 

al., 2000) 

Lopheucoila 

anastrepha 


(Gallardo & Diaz, 1999; Guimaraes et al., 1999) 

Aganaspis pelleranoi An Eucoilid parasite which seeks out larvae through fruit holes (Sivinski, et 

al., 1997). From Brasil (Guimaraes, et al., 1999). 

Dicerataspis flavipes An Eucoild larval parasite from Brasil (Guimaraes, et al., 1999). 

Doryctobracon 

areolatus 


(Souza-Filho, M.F., et al., 1999) 

Aganaspis 

An Eucoilid larval parasite from Brazil (Guimaraes, et al., 1999). 

nordlanderi 


al., 1997). From Brasil (Guimaraes, et al., 1999). 

Doryctobracon 

areolatus 

A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995; Lopez, et 

al., 1999). 

Nealiolus sp. A Braconid parasite from Mexico (Hernandez-Ortiz, 1994). 

Doryctobracon 

areolatus 

Doryctobracon 

areolatus 

Anastrepha bezzi Biosteres 

longicaudatus 


vindemmiae 

Anastrepha 

consobrina 

Anastrepha 

cordata 

Anastrepha 

crebra 

Anastrepha 

distincta 

A parasitic Braconid from N. and S. America, est. in Florida (Wharton & 

Gilstrap, 1983; Wharton & Marsh, 1978). 

Biosteres brasiliensis A parasitic Braconid from Brazil (Thompson, 1943). 

Doryctobracon 

areolatus 

A solitaryBraconid entoparasitoid from Brazil (Leonel, et al., 1995). 

A Braconid parasite from Bolivia (Rogg & Camacho, 2003). 

A Pteromalinid parasite from Bolivia 

A Braconid larval-pupal parasite of late instars from N. and S. America, est. 

in Florida (Wharton & Gilstrap, 1983; Wharton & Marsh, 1978). 

Opius hirtus A Braconid parasitefrom Mexico (Hernandez-Ortiz, 1994). 

Microcrasis sp. A Braconid parasite from Mexico (Hernandez-Ortiz, 1994). 

Utetes anastrephae A Braconid parasite from Mexico (Hernandez-Ortiz, 1994). 

Doryctobracon 

areolatus 

Doryctobracon 

crawfordi 

A Braconid larval-pupal parasite of late instars from Mexico (Hernandez-Ortiz, 

1994). 

A Braconid larval-pupal parasite, sensitive to heat and dryness, from Mexico 

to Northern S. America (Hernandez-Ortiz, 1994). 

Opius sp. A solitaryBraconid entoparasitoid from Brazil (Leonel, et al., 1995). 


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Anastrepha 

fraterculus 

Aganaspis pelleranoi A Eucoilid parasite from Venezuela (Katiyar, et al., 1995). 

Doryctobracon 

areolatus 

A Braconid larval-pupal parasite of late instars from N. and S. America (Jiron 

& Mexzon, 1989; Aguiar-Menezes, et al., 2001). 

Phaenocaroa sp. A Braconid parasite from Columbia (Carrejo & Gonzalez, 1999) 

Phaenocarpa 

A Braconid parasite from Columbia (Trostle, et al., 1999) 

pericarpa 

Biosteres brasiliensis A Braconid larval parasite from Brasil (Silvestri, 1939; Thompson, 1943; 

Leonel, et al., 1995; Narayanan & Chawla 1962; Zucchi, et al., 1999). 

Doryctabracon 

areolatus 

Doryctobracon 

crawfordi 

Doryctobracon 

trinidadensis 

Biosteres 

(=Diachasmimorpha) 

longicaudata 


(Sivinski, et al., 1997; Wharton & Gilstrap, 1983; Wharton & Marsh, 1978; 

Leonel, et al., 1995; Narayanan & Chawla 1962; Katiyar, et al., 1995; 

Hernandez-Ortiz, 1994; Zucchi, et al., 1999; Lopez, et al., 1999; 

Aguiar-Menezes, et al., 2001; Aguiar-Menezes & Menezes, 2002; Sivinski, et 

al., 2000: Aluja, et al., 2000; Ovruski, 2002; Marchiori, et al., 2000; 

Menezes & Menezes, 2001). 


to Northern S. America (Sivinski, et al., 1997; Katiyar, et al., 1995; 

(Hernandez-Ortiz, 1994; Lopez, et al., 1999; Sivinski, et al., 2000). 

A Braconid larval-pupal parasite from Peru, where it and D. areolatus exert 

17% of the natural biological control (Cruz, 1998). 

A widely distributed solitary Braconid late-instar larval parasite from the 

South Pacific now ranging from Mexico to C. and S. America (Sivinski, et al., 

1997;Lopez, et al., 1999; Sivinski, et al., 2000; Rogg & Camacho, 2003). 

Aganaspis pelleranoi An eucoilid parasite from Argentina, Venezuela which seeks out larvae 

through fruit holes (Sivinski, et al., 1997; Thompson, 1943; Narayanan & 

Chawla 1962; Katiyar, et al., 1995; Lopez, et al., 1999; Sivinski, et al., 

2000; Aguiar-Menezes, et al., 2001; Ovruski, 2002) From Brazil (Guimaraes, 

et al., 1999; Marchiori, et al., 2000). 

Bracanastrepha 

anastrephae ( 

=Utetes anastrephae) 

A Braconid solitary larval-pupal parasite from S. and C. America, the 

Carribean and Florida (Wharton &Gilstrap, 1983; Thompson, 1943; Leonel, 

et al., 1995; Zucchi, et al., 1999, Narayanan & Chawla 1962; Lopez, et al., 

1999; Aguiar-Menezes, et al., 2001; Aguiar-Menezes & Menezes, 2002; 

Sivinski, et al., 2000; Menezes & Menezes, 2001). 

Opius bellus A Braconid parasite from S. and C. America (Wharton & Gilstrap, 1983; 

Leonel, et al., 1995; Narayanan & Chawla 1962; Zucchi, et al., 1999; 

Aguiar-Menezes, et al., 2001; Aguiar-Menezes & Menezes, 2002; Menezes 

& Menezes, 2001). 

Doryctobracon A Braconid parasite in Brazil (Zucchi, et al., 1999; Leonel, et al., 1995). 

fluminensis 

Asobara sp. An Alysiid parasite in Brazil (Zucchi, et al., 1999). 

Biosteres areolatus A Braconid parasite in Brazil (Thompson, 1943). 


argentina 

A Braconid parasite from Argentina (Thompson, 1943; Narayanan & Chawla 

1962). 

Ganaspis carvalhoi A Cynipid parasite from Brazil (Thompson, 1943; (Narayanan & Chawla, 

1962).) 

Coeloides 

anastrephae 

A Braconid parasite from Argentina (Narayanan & Chawla 1962). 


PPQ


Anastrepha 

grandis 

Anastrepha 

interrupta 

Anastrepha 

leptozona 

Anastrepha 

ludens 



Pseudecoila 

brasiliensis 

A Cynipid parasite(Narayanan & Chawla 1962). 

Opius anastrephae A Braconid parasite from Venezuela (Katiyar,et al., 1995); from Mexico 

(Sivinski, et al., 2000). 

Idiasta sp. A Braconid parasite from Venezuela (Katiyar, et al., 1995). 

Lopheucoila sp. A Cynipid parasite from Mexico (Hernandez-Ortiz, 1994). 

Lopheucoila 

anastrephae 

Odontosema 

anastrephae 


(Gallardo & Diaz, 1999; Guimaraes, et al., 1999). 

A Eucoilid larval-pupal parasite from Mexico, Brazil (Guimaraes, et al., 1999; 

Lopez, et al., 1999; Aguiar-Menezes & Menezes, 2002). 

Odontosema sp A Eucoilid parasite from Mexico (Hernandez-Ortiz, 1994). 

Dirhinus (=Coptera) 

haywardi 


indica 

A Diapriinid pupal parasite from Argentina (Loiacono, 1981 (Lopez, et al., 

1999); from Mexico (Sivinski, et al., 2000). In Bolivia (Rogg & Camacho, 

2003). 

A Eulophid larval-pupal parasite from Mexico (Lopez, et al., 1999; Sivinski, et 

al., 2000). 

Spalangia endius A Pteromalid parasite from Brazil (Aguiar-Menezes & Menezes, 2002). 


vindemmiae 

Trichopria 

anastrephae 

Biosteres 

longicaudatus 


vindemmiae 


anastrephae 

Doryctobracon 

anastrephilum 

Doryctobracon 

areolatus 

A Pteromalid parasite from Brazil (Aguiar-Menezes & Menezes, 2002; 

Marchiori, et al., 2000); from Mexico (Sivinski, et al., 2000) In Argentina 

(Ovruski, 2002). In Bolivia (Rogg & Camacho, 2003). 

A Diapriid parasite from Brazil (Aguiar-Menezes, et al., 2001) 

A Braconid parasite recorded from Bolivia (Rogg & Camacho, 2003). 

A Pteromalid parasite in Bolivia (Rogg & Camacho, 2003). 

A Braconid larval parasite from Florida (Baranowski, et al., 1993). 

A Braconid larval parasite form Florida(Baranowski, et al., 1993; Wharton & 

Marsh, 1978). 

A solitary Braconid entoparasitoid from Brazil, Columbia (Leonel, et al., 

1995; Carrejo & Gonzalez, 1999; Menezes & Menezes, 2001). 

Opius sp. A solitary entoparasitoid from Brazil, Columbia (Leonel, et al., 1995; Carrejo 

& Gonzalez, 1999). 

Opius bellus A solitary entoparasitoid from Brazil ( Menezes & Menezes, 2001). 

Doryctobracon zeteki A solitary entoparasitoid from Columbia (Carrejo & Gonzalez, 1999). 

Aganaspis sp. An Eucoilid parasite from Columbia (Carrejo & Gonzalez, 1999). 

Utetes anastrephae A solitary entoparasitoid from Brazil ( Menezes & Menezes, 2001). 

Opius longicaudatus 

var. novocaledonicus 

ABraconid well adapted to this host (Clausen, 1956). 

Coptera hayardi A Diapriid pupal endoparasite from Latin America (Sivinski, et al., 1998; 

Lopez, et al., 1999; Sivinski, et al., 1998). 

Doryctabracon 

areolatus 


(Sivinski, et al., 1997; Wharton & Gilstrap, 1983; Aluja, et al., 1990; Lopez, 

et al., 1999; Sivinski, et al., 2000). 


PPQ




Anastrepha 

ludens (cont.) 

Anastrepha 

macrura 

Anastrepha 

manihoti 

Anastrepha 

mombinpraeopta 

ns 

Anastrepha 

montei 

Doryctobracon 

crawfordi 


vindemiae 

Diachasmimorpha 

longicaudata 


to Northern S. America (Sivinski, et al., 1997; Wharton & Gilstrap, 1983; 

Thompson, 1943; Narayanan & Chawla 1962; Hernandez-Ortiz, 1994; Aluja, 

et al., 1990Gonzalez-Hernandez & Tejada, 1979; Lopez, et al., 1999; 

Sivinski, et al., 2000). 

A Pteromalid pupal parasite from Mexico (Sivinski, et al., 2000). 


South Pacific now ranging from Mexico to and S. America (Sivinski, et al., 

1997; Aluja, et al., 1990; Lopez, et al., 1999; Sivinski, et al., 2000). 


al., 1997; Lopez, et al., 1999). 

Opius africanus A Braconid parasite from C. & S. America (Wharton & Gilstrap, 1983). 

Solenopsis geminata An efficient Formicid predator (94%) of pupal, prepupal stages & teneral 

emergent flies in Texas (Thomas, 1995). 

Chromocryptus 

mesorufus 

An Ichneumonid parasite from Mexico (Thompson, 1943; Narayanan & 

Chawla 1962). 

Cratospila ridibunda A Braconid parasite from Mexico (Thompson, 1943). 

Eucoila sp. A Cynipid possibly larval parasite from Mexico (Thompson, 1943; Baker, et 

al., 1944). 

Galesus sp. A Diapriid possibly pupal parasite from Mexico (Thompson, 1943; Baker, et 

al., 1944). 

Villa scylla A Bombyliid (Diptera) from Mexico (Thompson, 1943) 

Peromyscus leucopus A Cricetid mouse, a pupal predator, common in N.America, accounts for 34% 

(White-footed mouse) of pupae with species following (Thomas, 1993). 

Peromyscus boylii 

(Brush mouse) 

A Cricetid mouse, pupal predator in W. N. America (Thompson, 1993). 

Opius cereus A Braconid parasite from Mexico; Narayanan & Chawla 1962; 

Gonzalez-Hernandez & Tejada, 1979). 


anastrephae 

A Braconid parasite from Mexico (Aluja, et al., 1990). 

Ganaspis carvalhoi A non-specific Cypnipid parasite from Mexico (Aluja, et al., 1990). 

Aceratoneuromyia A larval-pupal Eulophid parasite from Mexico (Lopez, et al., 1999; Sivinski, et 

indica 

al., 2000). 

Homaeotarsus sp. A predaceous Staphylinid beetle (Thomas, 1995). 

Xenopygus analis A predaceous Staphylinid beetle (Baker, et al., 1944). 

Belomicjis rufipennis A predaceous Staphylinid beetle (Aluja, 1994). 

Utetes anastrephae A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995). 

Utetes anastrephae A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995). 

Doryctobracon 

areolatus 


in Florida (Wharton & Gilstrap, 1983). 


PPQ


Anastrepha 

montei (cont.) 

Anastrepha 

obliqua 



Opius bellus A Braconid parasite from S. and C. America (Wharton & Gilstrap, 1983; 

Leonel, et al. 1995). 

Spalangia 

A Pteromalid pupal parasite (Clausen, 1956) 

philippinensis 

Spalangia endius A Pteromalid parasite from Brazil (Aguiar-Menezes & Menezes, 2002). 

Doryctabracon 

areolatus 

Doryctobracon 

crawfordi 


longicaudata 

Utetes anastrephae 

(= Brachanastrepha 

anastrephae) 

A Braconid larval-pupal parasite of late instars from Mexico to Columbia to 

Argentina (Sivinski, et al., 1997;Wharton & Gilstrap, 1983; Leonel, et al., 

1995; Katiyar, et al., 1995; Jiron & Mexzon, 1989; Hernandez-Ortiz, et al., 

1994; Aluja, et al., 1990; Carrejo & Gonzalez, 1999; Lopez, et al., 1999; 

Aguiar-Menezes, et al., 2001; Aguiar-Menezes & Menezes, 2002; Sivinski, et 

al., 2000; Aluja, et al., 2000; Menezes & Menezes, 2001). 

A Braconid pupal predator, common in N.America, accounts for 34% of pupae 

with species following (Thomas, 1993; Lopez, et al., 1999). 


South Pacific now ranging from Mexico to S. America (Sivinski, et al., 1997; 

Aluja, et al., 1990; Lopez, et al., 1999; Sivinski, et al., 2000; Rogg & 

Camacho, 2003)). 

A solitary Braconid larval-pupal parasite from Florida to Argentina (Sivinski, et 

al., 1997; Wharton & Gilstrap, 1983; Leonel, et al., 1995; Hernandez-Ortiz, 

et al., 1994; Aluja, et al., 1990; Lopez, et al., 1999; Aguiar-Menezes, et al., 

2001; Aguiar-Menezes & Menezes, 2002; Sivinski, et al., 2000; Menezes & 

Menezes, 2001). 

Biosteres tryoni A Braconid egg-larval parasite from eastern Australia, est. Hawaii (Wharton & 


Coptera haywardi A diapriid endoparasitoid pupal parasite (Sivinski, et al., 2000). 

Opius bellus A Braconid parasite from S. and America (Wharton & Gilstrap, 1983; Leonel, 

et al., 1995; Katiyar, et a 1995; Aguiar-Menezes & Menezes, 2002; 

Menezes & Menezes, 2001). 

Asobara anastrephae A solitary Alysinne Braconid entoparasitoid from Brazil (Leonel, et al., 1995). 

Opius sp. A solitary Braconid entoparasitoid from Brazil, two species from Venezuela 

(Leonel, et al., 1995; Katiyar, et al., 1995). 

Doryctibracon sp. A Braconid parasite from Venezuela (Katiyar, et al., 1995). 

Opius anastrephae A Braconid parasite from Venezuela (Katiyar, et al., 1995), 

Aganaspis pelleranoi A Eucoilid parasite from Venezuela (Katiyar, et al., 1995; Lopez, et al., 1999), 

Brazil (Guimaraes et al., 1999; Aguiar-Menezes, et al., 2001; Aguiar-Menezes 

& Menezes, 2002). 

Eucoline cynipids Three unidentified species of Eucoiline cynipids from Costa Rica (Jiron & 

Mexzon, 1989). 


indica 

A Eulophid parasite from Costa Rica, Mexico (Jiron & Mexzon, 1989; Aluja, et 

al., 1990; Sivinski, et al., 2000). 

Trichopria sp. A Diapriid parasite from Costa Rica; Jiron & Mexzon, 1989). 

Opius hirtus A Braconid parasite from Mexico (Hernandez-Ortiz, et al., 1994). 

Odontosema 

anastrephae 

A Figitid parasite from Brazil (Aguiar-Menezes & Menezes, 2002). 

Pachycrepoideus A Pteromalid parasite from Brazil (Aguiar-Menezes & Menezes, 2002); from 

vindemmiae 

Mexico (Sivinski, et al., 2000). In Bolivia (Rogg & Camacho, 2003). 


PPQ




Anastrepha 

ocresia 

Anastrepha 

ornata 

Anastrepha 

parallela 

Anastrepha 

pickeli 

Anastrepha 

pseudoparallela 

Anastrepha 

serpentina 

Anastrepha 

serpentina 

(Cont.) 

Doryctobracon 

fluminensis 

Doryctobracon 

areolatus 

A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995). 


in Florida (Wharton & Gilstrap, 1983; Carrejo & Gonzalez, 1999). 

Doryctobracon sp. A Braconid parasite from Venezuela (Katiyar, et al., 1995). 

Eucoiline cynipids Three unidentified species of Eucoiline cynipids from Costa Rica (Jiron & 

Mexzon, 1989) 

Doryctobracon A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995). 

areolatus 


fluminensis 

Lopheucoila 

anastrephae 

Biosteres 

(Doryctobracon) 

areolatus 


(Gallardo & Diaz, 1999; Guimaraes, et al., 1999) 

A larval parasite from Brasil (Silvestri, 1939). A Braconid larval-pupal 

parasite of late instars from N. and S. America, est. in Florida (Wharton 

&Gilstrap, 1983; Leonel, et al., 1995; Katiyar, et al.,1995; Aluja, et al., 

1990; Sivinski, et al., 2000; Aguiar-Menezes, et al., 2001; Menezes & 


Coptera haywardi A Diapriid pupal parasite from Venezuela (Garcia & Montilla, 2001). 

Opius anastrephae A Braconid larval parasite from Brasil, Mexico (Silvestri, 1939; Aluja, et al., 

1990; Menezes & Menezes, 2001). 

Opius cereus A Braconid parasite from Trinidad (Thompson, 1943;Narayanan & Chawla 

1962). 

Opius trinidadensis A Braconid parasite from Trinidad (Thompson, 1943;Narayanan & Chawla 

1962). 


brasiliensis 

Opius bellus A solitary entoparasitoid from Brazil, Venezuela (Leonel, et al., 1995; Katiyar, 

et al., 1995; Menezes & Menezes, 2001). 

Ganaspis carvalhoi A Cynipid parasite from Brazil, Mexico (Narayanan & Chawla, 1962; Aluja, et 

al., 1990). 

Trichopria 

A Diapriid parasite from Brazil (Narayanan & Chawla 1962). 

anastrephae 


indica 

A Eulophid parasite from Venezuela (Katiyar, et al., 1995); from Mexico 


Aganaspis pellenaroi An Eucoilid larval parasitoid from Mexico (Sivinski, et al., 2000). 


longicaudata 

Doryctobracon 

crawfordi 


South Pacific now ranging from Mexico to N. and S. America (Jiron & Mexzon, 

1989; Aluja, et al., 1990; Sivinski, et al., 2000). 

A Braconid pupal predator, common in N. America (Aluja, et al., 1990; 



PPQ


Anastrepha 

sororcula 

Anastrepha 

spatulata 

Anastrepha 

striata 



Doryctobracon zeteki A solitary Braconid entoparasitoid from Columbia (Carrejo & Gonzalez, 

1999). 


vindemiae 

A Pteromalid pupal parasite from Mexico (exotic) (Lopez, et al., 1999; 


Utetes anastrephae A solitary Braconid entoparasitoid from Mexico (Sivinski, et al., 2000). 

Doryctobracon 

areolatus 

Doryctobracon 

brasiliensis 

A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995; 

Aguiar-Menezes & Menezes, 1997; Aguiar-Menezes, et al., 2001; 

Aguiar-Menezes & Menezes, 2002; Menezes & Menezes, 2001). 

A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995). 

Utetes anastrephae A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995; 

Aguiar-Menezes, et al., 2001; Aguiar-Menezes & Menezes, 1997; Menezes 

& Menezes, 2001). 

Odontosema 

A Figitid parasite from Brazil (Aguiar-Menezes & Menezes, 2002). 

anastrephae 

Opius bellus A Braconid parasite from Brazil (Aguiar-Menezes, et al., 2001; Menezes & 

Menezes, 2001) 

Aganaspis pelleranoi A Figitid parasite from Brazil (Aguiar-Menezes, et al., 2001; Aguiar-Menezes & 


Pachycrepoideus A Pteromalid parasite from Brazil (Aguiar-Menezes & Menezes, 2002). 

vindemmiae 

Doryctabracon 

areolatus 

Doryctobracon 

toxotrypanae 

Doryctabracon 

areolatus 

Doryctobracon 

crawfordi 


longicaudata 

(= Biosteres 

longicaudatus) 

Aganaspis 

nordlanderi 


South Pacific now ranging from Mexico to N. and S. America (Aluja, et al., 

2000) 

A larval-pupal host restricted Braconid parasite from Costa Rica, Mexico 

(Aluja, et al., 2000) 


South Pacific now ranging from Mexico to N. and S. America (Sivinski, et al., 

1997; Wharton & Gilstrap, 1983; Jiron & Mexzon, 1989; Aluja, et al., 1990; 

Lopez, et al., 1999; Sivinski, et al., 2000; Aluja, et al., 2000). 


to Northern S. America (Sivinski, et al., 1997; Wharton & Gilstrap, 1983; 

Thompson, 1943; Narayanan & Chawla 1962; Katiyar, et a; 1995; Jiron & 

Mexzon, 1989; Hernandez-Ortiz, 1994; Aluja, et al., 1990; Lopez, et al., 

1999; Sivinski, et al., 2000). 

A Braconid larval-pupal parasite of late instars from Mexico to Columbia to 

Argentina (Sivinski, et al., 1997; Leonel, et al., 1995; Katiyar, et al., 1995: 

Jiron & Mexzon, 1989; Hernandez-Ortiz, 1994; Carrejo & Gonzalez, 1999; 

Lopez, et al., 1999; Sivinski, et al., 2000; Rogg & Camacho, 2003). 

An Eucoilid larval parasite from Brazil (Guimaraes, et al., 1999). 


al., 1997). From Venezuela (Katiyar, et al., 1995;), Mexico (Lopez, et al., 

1999; Sivinski, et al., 2000). From Brasil (Guimaraes, et al., 1999). 

Fopius arisanus 

(=Biosteres arisanus) 

A Braconid larval-pupal parasite from Indo-Australia, now America, (Wharton 

& Gilstrap, 1983; Quimio & Walter, 2001) 

Opius bellus A Braconid parasite from N. and S. America (Wharton & Gilstrap, 1983). 


PPQ




Anastrepha 

suspensa 

Chromocryptus 

mesorufus 

An Ichneumonid parasite from Mexico (Thompson; 1943; Narayanan & 

Chawla (1962). 

Opius cereus A Braconid parasite from Trinidad (Thompson, 1943; (Narayanan & Chawla 

1962). 

Dorybctobracon 

(=Opius) trinidadensis 

A Braconid parasite from Trinidad (Thompson, 1943; (Narayanan & Chawla 

1962). In Peru and Bolivia (Rogg & Camacho, 2003). 

Doryctobracon zeteki A Braconid parasite from Venezuela, Columbia (Katiyar, et al., 1995; Carrejo 

& Gonzalez, 1999). 

Doryctobracon sp. A Braconid parasite from Venezuela (Katiyar, et al., 1995). 

Idiasta sp. A Braconid parasite from Venezuela (Katiyar, et al., 1995). 

Eucoiline cynipids Three unidentified species of Eucoiline cynipids from Costa Rica (Jiron & 

Mexzon, 1989). 

Trichoptera sp. A Diapriid parasite from Costa Rica (Jiron & Mexzon, 1989; Lopez, et al., 

1999). 


indica 

A Eulophid parasite from Costa Rica (Jiron & Mexzon, 1989). 

Lopheucoila sp. A Cynipid parasite from Mexico (Hernandez-Ortiz, 1994). 

Odontosema sp. A Cynipid parasite from Mexico (Hernandez-Ortiz, 1994). 

Utetes anastrephae A Braconid larval-pupal parasite from Mexico (Lopez, et al., 1999; Sivinski, et 

al., 2000). 

Odontosema 

anastrephae 

A Braconid larval-pupal parasite from Mexico (Lopez, et al., 1999; Sivinski, et 

al., 2000). 

Coptera haywardi A Diapriid pupal parasite from Venezuela (Garcia & Montilla, 2001). 


indica 


vindemiae 

Diachasmimorpphia 

longicaudata 

(= Biosteres 

longicaudatus) 

A Braconid larval-pupal parasite from Mexico (Lopez, et al., 1999; Sivinski, et 

al., 2000). 

A Pteromalid pupal parasite from Mexico (Sivinski, et al., 2000). In Bolivia 

(Rogg & Camacho, 2003). 

A Braconid late instar larval-pupal parasite from SE Asia which introduces a 

virus and a new protein into the host, preventing encapsulation of the 

parasite. The result is a 95% cut in fly populations if mass released. (Adams, 

1994; Investigator, 1999; Baranowski, et al., 1993; Wharton & Gilstrap, 

1983; Wharton & Marsh, 1978; Baranowski, et al., 1993; Sivinski, et al., 

1999). 

Coptera haywardi A Diapriid pupal endoparasitoid from Latin America, but unable to develop on 

irradiated hosts or parasitized hosts (Menezes, et al., 1998; Sivinski, et al., 

1998). Preference for this host 

Spalangia cameroni A nonspecific Pteromalid larval parasite from Florida (Baranowski, et al., 

1993). 

Spalangia endius A nonspecific Pteromalid larval parasite from Florida (Baranowski, et al., 

1993). 


vindemiae 

A nonspecific Pteromalid larval parasite from Florida (Baranowski, et al., 

1993). 

Cothonaspis sp. A nonspecific Eucoilid larval parasite from Florida (Baranowski, et al., 1993). 

Trichopria sp. A nonspecific Diapriid larval parasite from Florida (Baranowski, et al., 1993). 

Bracanastrepha A Braconid solitary larval-pupal parasite from S. and N. America, the 

anastrephae (= Caribbean and Florida (Baranowski, et al., 1993; Wharton & Gilstrap, 1983; 

Utetes amastrephae) Sivinski, et al, 1999). 


PPQ


Anastrepha 

suspensa (Cont.) 

Anomoia 

permunda 

Doryctibracin 

anastrephilus 



A Braconid larval parasite from Florida (Baranowski, et al., 1993; Wharton & 

Marsh, 1978). 

Opius concolor A Braconid parasite from northern Africa, intr. through the Mediterranean 

(Wharton & Gilstrap, 1983). 

A Braconid parasite (Narayanan & Chawla 1962). 

lceratoneauromyia A Eulophid larval parasite from Costa Rica (Baranowski, et al., 1993). 

indicum 

Biosteres 

A Braconid larval parasite from Hawaii (Baranowski, et al., 1993). 

vandenboschi 


tryoni 

Doryctobracon 

areolatus 

A Braconid larval parasite from Hawaii Origin: Queensland (Baranowski, et 

al., 1993). 

A Braconid larval parasite from Trinidad with parasitic rates of 49% 

(Baranowski, et al., 1993). A Braconid larval-pupal parasite of late instars 

from Mexico to Argentina Imported to Florida (Sivinski, et al., 1997; Sivinski, 

et al., 1999). 

Dirhinus himalayanus A Chaldid pupal parasite from Asia, also found on muscoid pupae (Sivinski, 

et al., 1998). 

Spalangia gemina A Pteromalid pupal parasite from Asia, also S. America (Sivinski, et al., 

1998). 

Doryctobracon 

trinidadensis 

A Braconid larval parasite from Trinidad (Baranowski, et al., 1993). 

Psyttalia concolor A Braconid larval parasite from France (Baranowski, et al., 1993). 

Psyttalia incisi A Braconid larval parasite from Hawaii (Baranowski, et al., 1993). 

Trybliographa daci An Eucoilid larval parasite from France (Baranowski, et al., 1993). 

Formicidae Four species of ants attacking larvae released on soil surface, causing up to 

100% mortality in Florida (Hennessey, 1998). 

Euborellia anulipes In Florida lab, this earwig (Carcinophoridae) was found eating larvae and 

pupae (Hennessey, 1998). 

Fopius arisanus In lab, this Braconid egg parasite from the Indo-Australian Region completed 

normal development (Lawrence, et al., 2000) 

Halticoptera laevigata A Pteromalid parasite from Russia, Europe, China, Japan (Kandybina, 1977). 

Bactrocera spp Bathyaulax sp. A Braconid parasite from India (Thompson, 1943) 

Bathyaulax 

trypaeniphaga 

A Braconid parasite form India (Thompson, 1943) 


albobalteata 

A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 


longicaudata 


Fopius arisanus A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

Fopius vandenboschi A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

Psyttalia incisi A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

Psyttalia makii A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

Utetes bianchii A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

Bactrocera 

aberrans 

Opius tryoni A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 


PPQ




Bactrocera 

albistrigata 

Bactrocera 

aquilonis 

Bactrocera 

arecae 

Bactrocera 

atrisetosa 

Bactrocera 

barringtoniae 

Bactrocera 

cacuminata 

Bactrocera 

cacuminata 

(Cont.) 


longicaudata 

A Braconid parasite from Malaysia and Thailand (Chinajariyawong, et al., 

2000). 

Fopius arisanus A Braconid parasite from Malaysia and Thailand (Chinajariyawong, et al., 

2000). 

Fopius vandenboschi A Braconid parasite from Malaysia and Thailand (Chinajariyawong, et al., 

2000). 

Dipterophagus daci A Dipterophagid (Strepsiptera) gregarious (2-6 per host) endoparasite from 

Australia which attacks the soft ternal adult stage,resulting in sterile hosts 

(Drew & Allwood, 1985). 

Diachasimiorpha 

longicaudata 




Psytttalia sp. nr 

fletcheri 


Biosteres deeralensis A Braconid larval parasite from Australia & New Guinea (Wharton & Gilstrap, 

1983; Cochereu, 1970). 


kraussii (=Biosteres 

kraussii) 

A Braconid larval parasite from Australia (Wharton & Gilstrap, 1983; 

Rungrojwanich & Walter, 2000). 

Opius fijiensis A Braconid egg-larval parasite from Australia, S. Pacific (Wharton & Gilstrap, 

1983). 

Fopius arisanus A Braconid egg-pupal parasite from the Australian Region (Quimio & Walter, 

2001). 

Biosteres arisanus A Braconid egg parasite from the Indo-Australian Region (Wharton & Gilstrap, 

1983). 


kraussii (= Biosteres 

kraussii) 

Biosteres 

vandenboschi 

A Braconid larval parasite from Australia (Wharton & Gilstrap, 1983; 

Snowball, 1966; Rungrojwanich & Walter, 2000). 

A Braconid1st stage larval Parasite from SE Asia, est. in Hawaii (Wharton & 

Gilstrap, 1983). 




Opius perkinsi A Braconid larval parasite from Australia (Wharton & Gilstrap, 1983). 

Opius longicaudatus An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, Australia, 

America, Carribean, Florida (Wharton & Gilstrap, 1983; Snowball, 1966). 


(=Opius oophilus) 

A Braconid egg-pupal parasite from the Australian Region (Quimio & Walter, 

2001). 

A Braconid egg-larval parasite from SE Asia & dominant in Hawaii. Females 

also carry bacteria and fungi that destroy eggs (Bosch & Haramoto, 1951; 

Clausen, 1956; Snowball, 1966). 


PPQ


Bactrocera 

calophylli 

Bactrocera 

carambolae 

Bactrocera 

carambolae - 

papayae 

Bactrocera 

caryeae 

Bactrocera 

caryeae (Cont.) 



Opius tryoni An Australian Braconid larval parasite present in small numbers (Snowball, et 

al., 1962; May & Kleinschmidt, 1954), 6.2% (Allman, 1939).Has a wide host 

range (Clausen, 1956; Silvestri, 1914; Wharton & Gilstrap, 1983; Snowball, 

1966). 

Bracon sp. A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 

Opius fletcheri A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 


longicaudata 



An internal solitary larval-pupal parasite from Malaysia (Ibrahim, et al., 

1994). From Thailand and Malaysia (Chinajariyawong, et al., 2000). 



Fopius vandenboschi A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 

Psyttalia incisi A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 

Psyttalia makii A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 

Psyttalia sp. nr 

fletcheri 

A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 

Psyttalia sp.nr makii A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 

The following 

parasites from 

(Vijaysegaran, 

1984).It is not 

possible to identify 

the fruit fly host, 

which is likely to be 

carambolae, based 

on location, but could 

be papayae or 

perhaps both of these 

species. 

Biosteres 

persulcatus 

A Braconid larval parasite from Malaysia causing over 75% parasitism with 

the species below (Vijaysegaran, 1984). 

Biosteres arisanus A Braconid larval parasite from Malaysia causing over 75% parasitism with 

the species above (Vijaysegaran, 1984). 

Biosteres oophilus A Braconid larval parasite from Malaysia causing negilible parasitism 

(Vijaysegaran, 1984). 

Biosteres 

longicaudatus 

A Braconid larval parasite from Malaysia causing 7.9% 

parasitism(Vijaysegaran, 1984). 

Opius fletcheri A Braconid egg-larval parasite from Malaysia causing 20.7% parasitism 


Opius manii A Braconid larval parasite from India (Fullaway, 1951; see White & 

Eston-Harris, 1992 for this and following entries). 

Opius compensans A Braconid parasite from India (Thompson, 1943; Narayanan & Chawla, 

1962). 

Opius incisi A Braconid parasite from India (Thompson, 1943; Narayanan & Chawla, 

1962). 


PPQ




Bactrocera 

caudata 

Bactrocera 

ciliatus 

Bactrocera 

correcta 

Bactrocera 

cucumis 

Opius persulcatus A Braconid parasite from India (Thompson, 1943; Narayanan & Chawla, 

1962). 

Opius oophilus A Braconid parasite from India (Narayanan & Chawla, 1962). 

Cothonaspis sp. A Braconid parasite from India (Narayanan & Chawla, 1962). 

Pseudeucoila A Braconid parasite from India (Narayanan & Chawla, 1962). 

Trybliographa daci A Braconid parasite from India (Narayanan & Chawla, 1962). 

Dirhinus giffardii A Braconid parasite from India (Narayanan & Chawla, 1962). 

Spalangia spp. A Pteromalid parasite from India (Narayanan & Chawla, 1962). 

Syntomosphyrum 

indicum 

An Eulophid parasite from India (Narayanan & Chawla, 1962). 

Trichopria sp. A Diapriid parasitefrom India (Narayanan & Chawla, 1962). 

Dirihinus giffardii A Chalcid pupal parasite from Pakistan (Ahmad, et al., 1975) 

Psyttalia incisi A Braconid egg-larval parasite from India (Kapoor, 1993). 

Galesus sp. A Diapriid parasite ffrom India (Kapoor, 1993). 

Pachycrepoideusvinde 

miae (= dubius) 

A Pteromalid parasite from India (Kapoor, 1993). 

Spalangia endius (= 

philippinensis) 

A Spalangiid parasite from India (Kapoor, 1993). 

Dirhinus sp. A Chalcid larval-pupal parasite from India (Narayanan & Chawla, 1962). 


longicaudata 



Fopius persulcatus A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 


Psyttalia fletcheri A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 


Psyttalia makii A Braconid parasite from Malaysia and Thailand (Chinajariyawong, et al., 

2000). 


fletcheri 


Psyttalia sp. nr makii A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

Utetes bianchii A Braconid parasite from Malaysia and Thailand (Chinajariyawong, et al., 

2000). 


PPQ


Bactrocera 

cucurbitae 

Bactrocera 

cucurbitae (cont) 

Psytallia fletcheri = 

Opius fletcheri 



Common Indian egg-larvalBraconid parasite, in Hawaii restricted to Melon Fly. 

Also in the Philippines, Guam, India and int. to Ryukyu Is. (Clausen, 1956; 

Wharton & Gilstrap, 1983; Thompson, 1943; Cochereau, 1970). Narayanan 

& Chawla, 1962, state that in the fall, this parasite attacks up to 20% of full 

grown larvae leaving the host, and develop in the pupa. Note that P. fletcheri 

prefers wild hosts for parasitization (Liquido, 1991; Vargas et al., 2000; 

Vargas, 2002). From Thailand (Chinajariyawong, et al., 2000). In Peru and 

Bolivia (Rogg & Camacho, 2003). 

Opius watersi Egg-larval Braconid parasite in North India & Ceylon (Clausen, 1956; 

Narayanan & Chawla, 1962). 

Opius angaleti Egg-larval Braconid parasite in Borneo (Clausen, 1956; Narayanan & Chawla, 

1962. 

Biosteres angaleti A Braconid larval-pupal parasite from the Indo-Australian Region (Wharton & 



(=Biosteres) dacusii 

A Braconid parasite from India (Wharton & Gilstrap, 1983). From Thailand 

(Chinajariyawong, et al., 2000) 

Biosteres hageni A Braconid parasite from Fiji reared in lab ( Wharton & Gilstrap, 1983). 

Diachasmimorpha (= 

Biosteres) 

longicaudatus 

= Psyttalia incisi (= 

Opius incisi) 

An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, Australia, 

America, Carribean, Florida (Wharton & Gilstrap, 1983; Wharton & Marsh, 

1978). From Thailand (Chinajariyawong, et al., 2000). 

A Braconid egg-larval parasite from SE Asia, est. in Hawaii, in lab (Wharton & 

Gilstrap, 1983). In India Kapoor, 1993). 

Diachasma fullawayi A Braconid parasite from Hawaii (Thompson, 1943). 

Diashasma tryoni A Braconid parasite from Hawaii (Thompson, 1943). 

Dirhinus giffardii A Chalcid pupal parasite from Hawaii (Thompson, 1943) Intro. to Reunion 

(Etienne, 1973; Narayanan & Chawla, 1962). From India (Kapoor, 1993). 

Fopius arisanus From Thailand and Malaysia (Chinajariyawong, et al., 2000). 

Galeus silvestrii A Diapriid parasite from Hawaii (Thompson, 1943). 

Ipobracon sp. A Braconid parasite from India (Thompson, 1943). 

Opius sp. A Braconid parasite from Ceylon (Thompson, 1943). 

Opius humilis A Braconid parasite from Hawaii (Thompson, 1943). 

Pachycrepoideus A Pteromalid parasite from Hawaii(Thompson, 1943; Narayanan & Chawla, 

dubius=vindemiae 1962). From India (Kapoor, 1993). 

Paranagrus osborni A Mymarid parasite from Hawaii(Thompson, 1943). 

Splangia cameroni A Pteromalid parasite from Hawaii(Thompson, 1943). 

Splangia hirta A Pteromalid parasite from Hawaii(Thompson, 1943). 

Tetrastichus 

giffardianus 

A Eulophid parasite from Hawaii (Thompson, 1943). 

Dirhinus anthracia A Chalcid parasite from S. India, imported to Ceylon, in Hawaii, Fiji. From 

=Dirhinus sp. ? Africa to Middle East, SE Asia, S. Pacific, Australia. Parasite of many fly 

families (Boucek & Narendran, 1981; Cochereau, 1970). 

Spalangia nigra A Pteromalid, pro larval-pupal parasite from Europe, N. America, pro 

imported into HawaiMany diverse hosts (Boucek, 1963). 

Spalangia cameroni A Pteromalid, pro larval-pupal parasite from Europe, Africa, Asia, & S. 

America, imported to Pacific Is. Many diverse hosts (Boucek, 1963). 


PPQ




Bactrocera 

curvipennis 

Bactrocera 

decipiens 

Bactrocera 

decurtans 

Bactrocera 

depressa 

Bactrocera 

distincta 

Bactrocera 

diversa 

Bactrocera 

dorsalis 

Pachyneuron 

vindemmiae 

Fopius (=Opius) 

vandenboschi 

A Pteromalid pupal parasite from Morocco (Thompson, 1943). Intro. Reunion 

(Etienne, 1973). 

A Braconid parasite from SE Asia (Cochereau, 1970). From Thailand 

(Chinajariyawong, et al., 2000). 

Spalangia sp. A Pteromalid parasite from SE Asia (Cochereau, 1970). 


indicum 

An Eulophid parasite from SE Asia (Cochereau, 1970). 

Spalangia 

philippinensis (= 

endius) 

A Pteromalid pupal parasite from Hawaii (Narayanan & Chawla, 1962). From 

India (Kapoor, 1993). 

Opius compensans A Braconid parasite from India with 10-12% parasitism (Narayanan & Chawla, 

1962). 

Dirhinus auratus A Chalcid parasite from Ceylon (Narayanan & Chawla, 1962). 

Dirhinus luzonensis A Chalcid parasite from India (Narayanan & Chawla, 1962). 

Spalangia afra A Pteromalid parasite (Narayanan & Chawla, 1962). 

Spalangia 



Spalangia 

stomoxysiae 


Philonthus sp. A Staphylinid predator from South Chin (Narayanan & Chawla, 1962). 

Opius # 4 A Braconid parasite from Borneo which prefers large 3rd instar larva 

(Marucci, 1952). 

Utetes bianchii From Thailand (Chinajariyawong, et al., 2000). 

Biosteres 

longicaudatus 


An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji,America, 

Carribean, Florida (Wharton & Gilstrap, 1983; Wharton & Marsh, 1978). 

From New Caledonia (Cochereau, 1970). 


1983). 




Diachasmimorpha An Indo-Australian Braconid 2-3rd stage larval parasite, now in Hawaii, Fiji, 

(=Opius, =Biosteres) America, Carribean, Florida % parasitism 1.8% (Wharton & Gilstrap, 1983; 

longicaudata 

Thompson, 1943; Wharton & Marsh, 1978; Fry, 1987; Narayanan & Chawla, 

1962; Serit & Tan, 1990; Ooi, 1984; Serit, et al., 1986; Kapoor, 1993; 

Vargas et al., 2000; Chinajariyawong, et al., 2000). 


albobalteata 

A Braconid from Thailand ( Chinajariyawong, et al., 2000). 


PPQ


Bactrocerus 

dorsalis (Cont.) 





A New Caledonian 2-3rd instar larval parasite ex. Dacus spp. in Hawaii 

(Clausen, 1956). 

var. malaiensis A 2-3rd instar larval parasite from Malaya, Borneo, India in Hawaii (Clausen, 

1956; Cochereau, 1970). 

var. taiensis A 2-3rd instar larval parasite ex. Dacus spp. from Thailand in Hawaii 

(Clausen, 1956; Cochereau, 1970) 

Note: This species seems to attack older larvae in fruit on the ground (Bess, 

et al., 1950) 

var. chocki From the Philippines (Cochereau, 1970). 

Opius compensans An egg-larval Braconid parasite from India, in Hawaii (Clausen, 1956). Small 

numbers only (Narayanan & Chawla, 1962; Narayanan & Chawla, 1962). 

Opius formosanus An egg-larvalBraconid parasite from Formosa in Hawaii (Clausen, 1956; 


Psyttalia fletcheri A Braconid parasite fromThailand ( Chinajariyawong, et al., 2000). 

Psyttalia (=Opius) 

incisi 

An egg-larvalBraconid parasite from SE Asia in Hawaii (Clausen, 1956; 

Wharton & Gilstrap, 1983; Cochereau, 1970; Narayanan & Chawla, 1962; 

Ooi, 1984; Vargas et al., 2000). In Thailand (Chinajariyawong, et al., 2000). 


Psyttalia sp. nr A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

fletcheri 


Opius kraussi An egg-larval Braconid parasite from Australia in Hawaii (Clausen, 1956). 

Fopius arisanus A Braconid egg/pupal parasite from Malaya in Hawaii, 4.7% parasitism; 24% 

in Malaysia (Clausen, 1956; Wharton & Gilstrap, 1983; Cochereau, 1970; 


Serit & Tan, 1990; Ooi, 1984; Serit, et al., 1986; Vargas et al., 2000). In 

Thailand (Chinajariyawong, et al., 2000). In Peru and 

Bolivia (Rogg & Camacho, 2003) 


A Braconid egg/pupal parasite which outcompetes other parasites (Wood, 

1998; from the Indo-Australian Region (Wharton & Gilstrap, 1983; Fry, 1987; 

Serit, et al., 1986). egg parasitism 17.2% (Serit & Tan, 1990; Liquido, 1991) 

Replaced Fopius vandenboschi (Vargas, 2002). 

Synonyms of the above (Quimio & Walter, 2001) 

Fopius (=Opius A Braconid 1st stage larval parasite from Malaya in Hawaii, 4.7% parasitism; 

=Biosteres) 

24% in Malaysia (Clausen, 1956; Wharton & Gilstrap, 1983; Cochereau, 

vandenboschi 

1970; Serit & Tan, 1990; Ooi, 1984; Serit, et al., 1986; Vargas et al., 2000). 

In Thailand (Chinajariyawong, et al., 2000). Replaced by Fopius arisanus 

(Vargus, 2002). 

Syntomosphyrum An Eulophid parasite ex. Dacus spp. from Malaya in Hawaii(Clausen, 1956; 

indicum 

Thompson, 1943). From SE Asia (Cochereau, 1970).In India at 66% total 

parasite pop., intr. Italy (Narayanan & Chawla, 1962). From India (Kapoor, 

1993). 

Tetrastichus dacicida An Eulophid pupal parasite ex. Dacus, Pardalaspis & Pterandrus spp. from 

Kenya in Hawaii(Clausen, 1956; Narayanan & Chawla, 1962) 


PPQ





persulcatus 

A primarily 1st to 2nd instar Braconid parasite from Malaya, India attacking 

larvae on fruit in trees (Bess, et al., 1950; Cochereau, 1970; Serit, et al., 

1986).With O. oophilus, is 23% of parasite pop. (Narayanan & Chawla, 

1962). In Thailand (Chinajariyawong, et al., 2000). 

Biosteres deeralensis A Braconid larval parasite from Australia & New Guinea reared in Hawaii 


Biosteres hageni A Braconid parasite from Fiji reared in lab (Wharton & Gilstrap, 1983). 


kraussii (=Biosteres 

kraussii) 

An Australian Braconid larval parasite (Wharton & Gilstrap, 1983). D. 

kraussii cannot successfully reproduce in Hawaiian B dorsalis 

(Rungrojwanich & Walter, 2000). 

Biosteres persulcatus A Braconid parasite from India, Malaysia parasitism .1% (Wharton & Gilstrap, 

1983; Thompson, 1943; Narayanan & Chawla, 1962; Serit & Tan, 1990). 

Biosteres skinneri A Braconid parasite from Malaysia, Philippines which more readily attacks 

Tephritids in cucurbits,reared in lab (Wharton & Gilstrap, 1983; Serit & Tan, 

1990). 

Biosteres tryoni An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, Australia, 

America, Carribean, Florida (Wharton & Gilstrap, 1983; Thompson, 1943; 

Wharton & Marsh, 1978; Fry, 1987). 

Opius fijiensis A Braconid egg-larval parasite from Australia, S. Pacific, in lab (Wharton & 


Opius makii A Braconid parasite from SE Asia (Wharton & Gilstrap, 1983; Thompson, 

1943; Cochereau, 1970; Narayanan & Chawla, 1962). 

Opius manii A Braconid parasite from India (Wharton & Gilstrap, 1983; Narayanan & 

Chawla, 1962; Kapoor, 1993). 

Opius mitis A Braconid parasite in Peru and Bolivia (Rogg & Camacho, 2003). 

Biosteres javanus A Braconid parasite from Java (Thompson, 1943; Narayanan & Chawla, 

1962). 

Galesus sp. A Diapriid parasite from Malaya, India (Thompson, 1943; Narayanan & 

Chawla, 1962; Kapoor, 1993). 

Opius fletcheri A Braconid parasite from Malaysia, India (Thompson, 1943; Narayanan & 

Chawla, 1962). 

Spalangia sp. A Pteromalid parasite from Malaysia (Thompson, 1943; Cochereau, 

1970;Narayanan & Chawla, 1962). 

Tachinaephagus sp. An Encyrtid from Malaysia (Thompson, 1943; Narayanan & Chawla, 1962). 

Tetrastichus sp. An Eulophid parasite from Malaysia (Thompson, 1943). 

Spalangia endius A Pteromalid larval-pupal (?) parasite with many hosts, world-dist (Boucek, 

1963). 

Spalangia gemina A Pteromalid pupal parasite with a wide host range from Asia through S. 

America (Boucek, 1963). 

Pachyneuron 

A Pteromalid parasite from SE Asia (Cochereau, 1970). 

vindemmiae 

Dirhinus giffardii A Chalcidid pupal parasite (Narayanan & Chawla, 1962). 

Cothonaspis sp. A Cynipid parasite (Narayanan & Chawla, 1962; Kapoor, 1993). 

Trybliographa daci A Cynipid parasite (Narayanan & Chawla, 1962. 

Pilinothrix sp. A Cynipid parasite from Malaya (Narayanan & Chawla, 1962). 

Dirhinus luzonensis A Chalcid parasite from Malaya (Narayanan & Chawla, 1962). 


PPQ


Bactrocera 

dorsalis (Cont.) 

Bactocera 

endiandrae 


dubius = vindemiae 



A Cynipid pupal parasite ffrom the Philippines (Narayanan & Chawla, 1962). 

From India (Kapoor, 1993). 

Spalangiaafra A Pteromalid parasite (Narayanan & Chawla, 1962). 

Spalangia grotiusi A Pteromalid parasite (Narayanan & Chawla, 1962). 

Spalangia 

A Pteromalid parasite (Narayanan & Chawla, 1962). From India (Kapoor, 


(=endius) 

1993). 

Spalangia simplex A Pteromalid parasite Narayanan & Chawla, 1962). 

Spalangia stomoxysia A Pteromalid parasite Narayanan & Chawla, 1962). 

Trichopria sp. A Diapriid parasite from India (Narayanan & Chawla, 1962). 

Thyreocephalus 

albertisi 

A Staphylinid predator from the Philippines (Narayanan & Chawla, 1962). 

Spalangia grotiusi A Pteromalid parasite Narayanan & Chawla, 1962). 

Spalangia 


(=endius) 

A Pteromalid parasite Narayanan & Chawla, 1962). From India (Kapoor, 

1993). 

Spalangia simplex A Pteromalid parasite Narayanan & Chawla, 1962). 

Spalangia stomoxysia A Pteromalid parasite Narayanan & Chawla, 1962). 

Trichopria sp. A Diapriid parasite from India (Narayanan & Chawla, 1962). 

Thyreocephalus 

albertisi 

A Staphylinid predator from the Philippines (Narayanan & Chawla, 1962). 

Anisolabis eteronoma An earwig (Carcinophiridae) predator which feeds on larvae in fruit and soil 

(Narayanan & Chawla, 1962). 

Sphingolabis 

An earwig (Labiidae) predator which feeds on larvae (Narayanan & Chawla, 

hawaiiensis 

1962). 

Dirhinus auratus A Chalcid parasite from India (Narayanan & Chawla, 1962; Kapoor, 1993). 


indica 

An Eulophid parasite from Malaysia (Ooi, 1984). 

Dirhinus anthracina A Chalcid parasite from India (Kapoor, 1993). 

Pseudeucoila sp. A Cynipid parasite from India (Kapoor, 1993). 

Syntomosphyrum afra An Eulophid parasite from India (Kapoor, 1993). 


grotiusi 

An Eulophid parasite from India (Kapoor, 1993). 


stomyoxysine 

An Eulophid parasite from India (Kapoor, 1993). 

Anisolabis eteronoma An earwig (Carcinophiridae) predator which feeds on larvae in fruit and soil 


Coptera silvestrii A Psiline parasite from Africa (Benin, Ghana, Niger, Nigeria, Senegal, Kenya, 

Monzambique, Zululand), introduced to Hawaii (Yoder & Wharton, 2002). 


Opius sp. ? africanus A Braconid larval parasite from Australia (?) (May & Kleinschmidt, 1954). 

Opius deeralensis A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 



PPQ




Bactrocera 

facialis 

Bactrocera 

fagraea 

Bactrocera 

frauenfeldi 

Bactrocera 

garciniae 

Bactrocera 

halfordiae 

Bactrocera 

halfordiae 

(Cont.) 

Bactrocera 

incisa 

Bactrocera 

irvingiae 

Bactrocera 

isolata 

Bactrocera 

jarvisi 

Biosteres deerlensis A Braconid larval parasite from Australia & New Guinea (Wharton & Gilstrap, 

1983). 


1983). 

Opius deeralensis A Braconid larval parasite from Australia (May & Kleinschmidt, 1954; 

Wharton & Gilstrap, 1983). 

Opius fletcheri A Braconid larval parasite from Australia (May & Kleinschmidt, 1954) 

Opius froggatti A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 

Opius perkinsi A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 

Opius fijiensi A Braconid egg-larvalparasite from Australia, S. Pacific (Wharton & Gilstrap, 

1983). 


1983; Cochereau, 1970). 

Biosteres 

longicaudatus 

An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, America, 



1983; Cochereau, 1970). 


longicaudata 



Opius tryoni An Australian Braconid larval parasite found in 52% of larvae in wild fruit 

(Gurney, 1910; May & Kleinschmidt, 1954). 


Opius kraussi A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 

Fopius schlingeri A egg-pupal Braconid parasite from Australia (Wharton, 1999) 

Biosteres 


longicaudatus Carribean, Florida (Wharton & Gilstrap, 1983; Wharton & Marsh, 1978). 

Dirhinus luzonensis A Chalcid parasite from the Philippines (Narayanan & Chawla, 1962). 


dubius 



longicaudata 

A Braconid larval parasite from Thailand (Chinajariyawong, et al., 2000). 



longicaudata 

From Thailand (Chinajariyawong, et al., 2000). 

Fopius arisanus From Thailand (Chinajariyawong, et al., 2000) 

Psyttalia fletcheri From Thailand (Chinajariyawong, et al., 2000) 

Opius flecheri A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 


PPQ


Bactrocera 

kanchanaburi 

Bactrocera 

kandiensis 

Bactrocera kirki 

Bactrocera 

kraussi 

Bactrocera 

kraussi (Cont.) 


kraussii 

(= Biosteres kraussii) 

(= Opius kraussii) 



A Braconid larval parasite from Australia (May & Kleinschmidt, 1954; 

Wharton& Gilstrap, 1983; Cochereau, 1970; Rungrojwanich & Walter, 2000). 


1983; Cochereau, 1970). 


1983). 

Opius perkinsi A Braconid larval parasite from Australia (Wharton & Gilstrap, 1983; 

Cochereau, 1970). 


(?) 



An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, Australia, 

America, Carribean, Florida (Wharton & Gilstrap, 1983). 

A Braconid parasite from SE Asia intr. to Australia (Cochereau, 1970). 

Spalangia sp. 


2001). 

A Pteromalid parasite from Australia (Cochereau, 1970). 

Trybliographa daci A parasite from Australia (Cochereau, 1970). 

Opius tryoni A Braconid parasite from Australia (Jarvis, 1925). 


longicaudata 









kraussii 

An Australian Braconid larval parasite. (Rungrojwanich & Walter, 2000; 

Wharton & Gilstrap, 1983; Cochereau, 1970). 


1983; Cochereau, 1970). 

Opius froggatti A Braconid parasite from Australia, New Caledonia (Wharton & Gilstrap, 

1983). 



Fopius arisanus A Braconid parasite from SE Asia intr. to Australia (Cochereau, 1970) 



2001). 


PPQ




Bactrocera lata Diachasmimorpha 

longicaudata 

Bactocera 

laticaudus 

Bactrocera 

latifrons 

A Braconid parasite from Malaysia (Chinajariyawong, et al., 2000). 

Opius perkinsi A Braconid larval parasite from Australia (Fullaway, 1950; May & 

Kleinschmidt, 1954; Wharton & Gilstrap, 1983; Narayanan & Chawla, 1962). 

Opius froggatti A Braconid larval parasite in Australia, New Caledonia described from 

numerous specimens (Fullaway, 1950; Wharton & Gilstrap, 1983; 

Cochereau, 1970; Narayanan & Chawla, 1962). 


Biosteres deeralensis A Braconid larval parasite from Australia (Wharton & Gilstrap, 1983; 



1983). 

Biosteres arisanus A Braconid egg parasite from the Indo-Australian Region (Wharton & Gilstrap, 

1983; Liquido, et al., 1994; Kapoor, 1993). 


(=Biosteres) 

longicaudatus 


Carribean, Florida (Wharton & Gilstrap, 1983; Wharton & Marsh, 1978; 

Liquido, et al., 1994). In Thailand (Chinajariyawong, et al., 2000). 

Biosteres persulcatus A Braconid parasite from India (Wharton & Gilstrap, 1983; Narayanan & 

Chawla, 1962; Liquido, et al., 1994; Kapoor, 1993). 

Fopius (=Biosteres) 

vandenboschi 


Gilstrap, 1983; Liquido, et al., 1994). From Thailand (Chinajariyawong, et 

al., 2000). 

Opius incisi A Braconid egg-larval parasite from SE Asia, est. in Hawaii (Wharton & 

Gilstrap, 1983; Liquido, et al., 1994; Kapoor, 1993). 



A Braconid larval-pupal parasite from India (Narayanan & Chawla, 1962). 

From Thailand (Chinajariyawong, et al., 2000). 


2001). 

Fopius deeralensis A Braconid parasite from Thailand (Chinajariyawong, et al., 2000) 

Dirhinus giffardii A Chalcid parasite from India (Narayanan & Chawla, 1962; Liquido, et al., 

1994). 

Dirhinus luzonensis A Chalcid parasite from the Philippines, India (Narayanan & Chawla, 1962; 

Liquido, et al., 1994). 


dubius 

A Pteromalid parasite from South China (Narayanan & Chawla, 1962; 

Liquido, et al., 1994). 




Psyttalia sp. nr A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

fletcheri 

2000). 


Dichasmimorpha An Opiine larval parasite from Australia, introduced to Hawaii for control 

kraussi 

(Duan & Messing, 2000; Messing, et al., 2000). 



PPQ


Bactrocera 

limbifera 

Biosteres 

longicaudatus 





Hardy, 1974). 

Biosteres skinneri A Braconid parasite from the Philippines which more readily attacks 

Tephritids in Cucurbits (Wharton & Gilstrap, 1983; Hardy, 1974). 

Opius makii A SEAsian Braconid parasite from the Philippines (Cochereau, 1970). 

Opius fletcheri A Braconid parasite from the Philippines (Hardy, 1974). 


dubius 

A Braconid parasite from the Philippines (Hardy, 1974). 

Galesus sp. A Diapriid parasite from the Philippines (Hardy, 1974). 

Spalangia sp. A Pteromalid parasite from the Philippines (Hardy, 1974). 

Bactrocera mayi 

Bactrocera 

melanota 




Bactrocera 

melas 


Opius kraussi A Braconid larval parasite from Australia (May & Kleinschmidt, 1954). 


Bactrocera 

minax 

Bactrocera 

murrayi 

Bactrocera 

musae 

Bactrocera 

mutabilis 



Diachasmimorpha A Braconid larval parasite from Bhutan (Van Achterberg, 1999) 

feijeni 


kraussii (= Opius 

kraussi (= Biosteres 

kraussii) 

An Australian Braconid larval parasite. (Rungrojwanich & Walter, 2000; May 

& Kleinschmidt, 1954; Wharton & Gilstrap, 1983). 


1983). 



Chelisoches morio An earwig (Chelisochidae) predator from Papua New Guinea preying on larvae 

(Greve & Ismay, 1983). Adults readily attack and consume 2-3 larvae per day 

(Smith, 1977). 


1983). 


1983). 

Opius oophilus A Braconid egg parasite from SE Asia & dominant in Hawaii, est. in Australia. 

Females also carry bacteria and fungi that destroy eggs (Bosch & Haramoto, 

1951; Clausen, 1956; Snowball, 1966). 

Dipterophagus daci A Dipterophagid (Strepsiptera) gregaruiys (2-6 per host) endoparasite from 

Australia which attacks the soft ternal adult stage, resulting in sterile hosts 

(Drew & Allwood, 1985) 



PPQ




Bactrocera 

neohumeralis 


Opius kraussi A Braconid larval parasite from Australia (May & Kleinschmidt, 1954; 

Snowball, 1966). 

Opius tryoni A Braconid larval parasite from Australia (May & Kleinschmidt, 1954; 

Snowball, 1966). 

Fopius schlingeri An egg-pupal Braconid parasite from Australia (Wharton, 1999) 



Bactrocera 

occipitalis 

Bactrocera oleae Psyttalia (=Opius) 

concolor 

Bactrocera oleae 

(Cont.) 


2001). 



(Drew & Allwood, 1985).. 

A Braconid larval parasite with 30% parasitism from N. Africa, S. Europe 

(Silvestri, 1914; Clausen, 1956; Wharton & Gilstrap, 1983; Thompson, 

1943; Fry, 1987; Narayanan & Chawla, 1962; Kapatos, et al., 1977; 

Nuenschwander, et al., 1983; Kimani-Nojou, et al., 2000; El-Heneidy, et al., 

2001). Introduced to Italy in 1933 for olive fly control (Kimani-Nojou, et al., 

2000). From Spain-- In Peru and Bolivia (Rogg & Camacho, 2003). 

Opius dacicida A Braconid larval parasite from Ethiopia, Italy, S. Africa (Silvestri, 1914; 

Wharton & Gilstrap, 1983; Thompson, 1943; Narayanan & Chawla, 1962). 

Opius lonsburyi A Braconid larval parasite from the Transvaal, Kenya (Silvestri, 1914; 


Opius africanus A Braconid larval parasite from South Africa, also in Italy (Silvestri, 1914; 


Sigalphus daci A Braconid parasite from South Africa, Ethiopia (Silvestri, 1914; Thompson, 

1943; Narayanan & Chawla, 1962). 

Bracon celer An important Braconid parasite from South Africa (Silvestri, 1914; Narayanan 

& Chawla, 1962). 

Galesus silvestrii A Proctotrupid pupal parasite from Nigeria bred in lab (Silvestri, 1914; 

Thompson, 1943; Narayanan & Chawla, 1962). 

Biosteres 

longicaudatus 

Achrysocharis 

formosa erythrea 

Achrysocharis 

formosa meridionalis 

Allocerellus 

inquirendus 

Euderus (Allomphale) 

cavasolae 

Amblymerus 

modestus 

Atoposoma 

variegatum 

Cratotrechus 

larvarum 


Carribean, Florida (Wharton & Gilstrap, 1983). 

An Eulophid parasite from Ethiopia (Thompson, 1943). 

An Eulophid parasite from S. Africa attacks small larvae (Thompson, 1943; 


An Encyrtid parasite from Ethiopia (Thompson, 1943). 

An Eulophid ectoparasite from Italy (Thompson, 1943). 

A Pteromalid parasite from Ethiopia (Thompson, 1943). 

An Eulophid parasite from Ethiopia (Thompson, 1943). 

An Eulophid parasite from Italy (Thompson, 1943). 


PPQ


Bactrocera oleae 

(cont) 



Dinarmus dacicida 

virescens 

A Miscogasterid parasite from Italy (Thompson, 1943). 

Dirhinus giffardii A Chalcid parasite from Italy (Thompson, 1943; Narayanan & Chawla, 1962). 

Ephialtes divinator A Ichneumonid from the Maritime Alps (Thompson, 1943). 

Eulophus sp. An Eulophid parasite from Libya (Thompson, 1943). 

Pnigalo (Eulophus) 

longulus 

An Eulophid parasite from France, Greece, Italy (Thompson, 1943; Narayanan 

& Chawla, 1962). 

Fopius arisanus A Braconid egg-pupal parasite. In the lab in the Mediterranean it was 

successfully reared using olive fly eggs as a host (Calvitti, et al., 2002). 

Macroneura sp. An Eupelmid parasite from Egypt (El-Heneidy, et al., 2001). 

Eulophus 

An Eulophid parasite from France, Italy (Thompson, 1943). 

pectinicornis 

Eupelmus sp. An Eupelmid parasite from Libya (Thompson, 1943) From Egypt (El-Heneidy, 

et al., 2001). 

Eupelmus afer An Eupelmid parasite from Ethiopia (Thompson, 1943; Narayanan & Chawla, 

1962). 

Eupelmus degeeri An Eupelmid parasite from Italy (Thompson, 1943). 

Eupelmus urozonus An Eupelmid parasite from Italy, France, Spain, Palestine (Thompson, 1943; 

Fry, 1987; Narayanan & Chawla, 1962; Nuenschwander, et al., 1983). 

Eurytoma sp. A Eurytomid parasite from Libya (Thompson, 1943) From Egypt (El-Heneidy, et 

al., 2001). 

Eurytoma aethiops A Eurytomid parasite from Italy (Thompson, 1943) 

Eurytoma rosae An Eurytomid parasite from France, Greece (Thompson, 1943; Narayanan & 


Eurytoma rufipes An Eurytomid parasite from Italy (Thompson, 1943). 

Metriocharis 

atrocyanea 

An Eurytomid parasite from Ethiopia (Thompson, 1943; Narayanan & Chawla, 

1962). 

Metriocharis viridis An Eurytomid parasite from Ethiopia (Thompson, 1943; Narayanan & Chawla, 

1962). 

Microbracon celer A Braconid parasite from Italy, Ethiopia, S. Africa (Thompson, 1943). 

Opius africanus 

orientalis 

A Braconid parasite from Ethiopia (Thompson, 1943; Narayanan & Chawla, 

1962). 

Dinarmus dacicida A Pteromalid parasite from Europe (Thompson, 1943; Narayanan & Chawla, 

1962). 

Opius ponerophagus A Braconid parasite from India (Thompson, 1943; Narayanan & Chawla, 

1962). 

Opius siculus A Braconid parasite from Sicily, 30% parasitism (Thompson, 1943; 

Narayanan & Chawla, 1962; Kapatos, et al., 1977). 

Pteromalus sp. A Pteromalid parasite from France (Thompson, 1943). 

Teleopterus notandus A Eulophid egg-larval parasite from Ethiopia (Thompson,1943; Narayanan & 


Tetrastichus gentilei A Eulophid parasite from Italy (Thompson,1943). 

Tetrastichus maculifer A Eulophid parasite from Ethiopia (Thompson,1943). 

Trichomalus 

spiracularis 

A Pteromalid parasite from Italy (Thompson, 1943). 


PPQ




Bactrocera 

osbeckiae 

Bactrocera 

pallida 

Pnigalio agraules A Eulopid parasite from Greece (Fry, 1987) From Egypt (El-Heneidy, et al., 

2001). 

Halticoptera daci A Pteromalid parasite from Africa (Narayanan & Chawla, 1962). 

Eurytoma (= 

Eupelmus) martelli 

Pnigalo 

mediterraneus 

An Eupelmid parasite from North Africa (Narayanan & Chawla, 1962; 

Nuenschwander, et al., 1983) From Egypt (El-Heneidy, et al., 2001). 

An Eulophid parasite from the Mediterranean area (Narayanan & Chawla, 

1962; Nuenschwander, et al., 1983). 

Lasioptera brevicornis A Cecidomyiid fly (Narayanan & Chawla, 1962. 

Lasioptera 

(Prolasioptera) 

berlesiana 

Belonuchus 

rufipennis 

A Cecidomyiid fly egg predator, associated with the fungus Camarosporium = 

Macrophoma dalmatica; maximum mortality of eggs30-50%(Narayanan & 

Chawla, 1962; Nuenschwander, et a1983). 

A Staphylinid larval predator intr. to Italy from Brazil (Narayanan & Chawla, 

1962). 

Carabus banoni A carabid whose larvae and adults are a larval-pupal predator in Crete 

(Nuenschwander, et al., 1983). 

Poecilus cupreus A carabid whose larvae are a larval-pupal predator in Crete (Nuenschwander, 

et al., 1983). 

Pterostichus creticus A carabid whose larvae are a larval-pupal predator in Crete (Nuenschwander, 

et al., 1983). 

Ocypus fulvipennis A staphlinid whose adults are a larval-pupal predator in Crete 


Ocypus olens A staphlinid whose adults are a larval-pupal predator in Crete 


Aphaenogaster 

simonelli 

Crematogaster 

sordidula 

Tetramorium 

caespitum 

An ant (Formicidae), capable of destroying both larvae and adults of olive fly 






Cyrtoptyx latipes A Pteromalid parasite from Crete (Nuenschwander, et al., 1983) from Egypt 

(El-Heneidy, et al., 2001). 

Cyttoptyx sp. A Pteromalid parasite from Egypt (El-Heneidy, et al., 2001). 




longicaudata 


Fopius persulcatus A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 


Biosteres deeralensis A Braconid larval parasite from Australia and New Guinea (Wharton & 




kraussii) 

An Australian Braconid larval parasite (Rungrojwanich & Walter, 2000; 


Opius fijiensis A Braconid egg-larval parasite from Australia, S. Pacific (Wharton & Glistrap, 

1983) 


PPQ


Bactrocera 

papayae - 

carambolae 

Bactrocera 

papayae 

Bactrocera 

passiflorae 



The following parasites from (Vijaysegaran, 1984). It is not possible to identiify the fruit fly hosts, 

which is eitherpapayae orcarambolae or perhaps both of these species. 

Biosteres persulcatus A Braconid larval parasite from Malaysia causing over 75% parasitism with 

the species below (Vijaysegaran, 1984). 

Biosteres arisanus A Braconid larval parasite from Malaysia causing over 75% parasitism with 

the species above (Vijaysegaran, 1984). 

Biosteres oophilus A Braconid larval parasite from Malaysia causing negilible parasitism 

(Vijaysegaran, 1984) 

Biosteres 

longicaudatus 

A Braconid larval parasite from Malaysia causing 7.9% 

parasitism(Vijaysegaran, 1984). 

Opius fletcheri A Braconid egg-larval parasite from Malaysia causing 20.7% parasitism 


Biosteres javanus A Braconid pupal parasite from Java (Dammerman, 1929). 


longicaudata 


vandenboschi 




2000). 

A Braconid egg-larval parasite from Malaysia (Chuan, et al., 1981). From 

Thailand and Malaysia (Chinajariyawong, et al., 2000). 



Psyttalia incisi A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 

Psyttalia makii A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 


fletcheri 


2000). 

Psyttalia sp. nr makii A Braconid parasite from Thailand and Malaysia (Chinajariyawong, et al., 

2000). 

Tetrastichus 

giffardianus 


indicum 


(=Opius oophilus) (= 

Biosteres arisanus) 

A Eulophidpupal parasite, orignially from West Africa, imported from India/ 

Australia, in Fiji, Hawaii (Simmons, 1935; Clausen, 1956; Thompson, 1943; 


An Eulophid pupal parasite from Fiji/Australia, in Samoa(Clausen, 

1956).From SE Asia, intr. Fiji from Australia (Cochereau, 1970; Narayanan & 


A Braconid egg parasite from the Indo-Australian Region (Wharton & Gilstrap, 

1983). 


2001). 

A Braconid larval parasite from SE Asia intr. to Australia, Fiji (Cochereau, 

1970) 

Biosteres hageni A Braconid parasite from Fiji ( Wharton & Gilstrap, 1983; Thompson, 1943; 



Gilstrap, 1983; Wharton & Marsh, 1978; Narayanan & Chawla, 1962). 

Opius fijiensis A Braconid egg-larval parasite from Australia, S. Pacific; parasitism 12.5% 

(Wharton & Gilstrap, 1983; Thompson, 1943; Cochereau, 1970; Narayanan 

& Chawla, 1962). 


PPQ




Bactocera 

passiflorae 

(Cont.) 

Bactrocera 

pedestris 

Bactrocera 

peninsularis 

Bactrocera 

pendleburi 

Bactrocera 


Bactrocera 

propinqua 

Opius humilus A Braconid larval parasite from South Africa, est. in Hawaii, Bermuda, Fiji 

(Wharton & Gilstrap, 1983; Narayanan & Chawla, 1962; Narayanan & 


Opius longicaudatus A Braconid larval parasite from SE Asia intr. to Fiji (Cochereau, 1970). 

var. malaiensis 

Spalangia endius A Pteromalid larval-pupal (?) parasite with many hosts, world-dist (Cochereau, 

1970). 

Phaenocarpa leveri A Braconid parasite from Viti Levu and Taveuni Islands (Narayanan & Chawla, 

1962). 


dubius 

A Pteromalid larval-pupal parasite from Fiji (Narayanan & Chawla, 1962). 

Pachyneuron sp. A Pteromalid parasite (Narayanan & Chawla, 1962). 

Spalangia cameroni A Pteromalid larval-pupal parasite (Narayanan & Chawla, 1962). 

Spalangia sp. A Pteromalid parasite (Narayanan & Chawla, 1962). 

Germalus pacificus A Lygaeid egg predator, most important in reducing populations (Narayanan & 

Chawla, 1962; Simmons, 1935). 

Opius fletcheri A Braconid parasite (Narayanan & Chawla, 1962). 

Biosteres 

longicaudatus 


Carribean, Florida(Wharton & Gilstrap, 1983; Wharton & Marsh, 1978). 

Biosteres skinneri A Braconid parasite from the Philippines which more readily attacks tephrids 

in Cucurbits (Wharton & Gilstrap, 1983). 

Biosteres 

vandenboschi 



Fopius schlingeri A egg-pupal Braconid parasite from Australia (Wharton, 1999). 




Diachasmimorpha A braconid parasite From Thailand (Chinajariyawong, et al., 2000). 

longicaudata 


longicaudata 

Bactrocera psidii Biosteres 

longicaudatus 

Bactrocera 

pyrifoliae 




var. novocaledonicus From New Caledonia (Cochereau, 1970). 


1983; Cochereau, 1970). 

Opius froggatti A Braconid parasite from Australia, New Caledonia (Wharton & Gilstrap, 

1983; Cochereau, 1970). 

Spalangia endius A Pteromalid larval-pupal (?) parasite with many hosts, world-dist (Cochereau, 

1970). 

Diachasmimorpha A Braconid parasite from Thailand (Chinajariyawong, et al., 2000). 

longicaudata 



PPQ






Bactrocera 

raiensis 


Bactrocera 

rufofuscula 

Fopius schlingeri A egg-pupal Braconid parasite from Australia (Wharton, 1999). 

Bactocera 

scutellaris 



Bactocera 

scutellaris 

Psyttalia fletcheri A braconid parasite from Thailand (Chinajariyawong, et al., 2000) 

Bactocera 

signatifera 

Bracon sp. A Braconid larval parasite from Australia(May & Kleinschmidt, 1954). 

Bactrocera 

strigifinis 

Biosteres sp. A Braconid larval parasite from Papua New Guinea (Greve & Ismay, 1983). 

Bactrocera tau Diachasmimorpha (= An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, America, 

Biosteres) 

Carribean, Florida (Wharton & Gilstrap, 1983; see also, White & 

longicaudatus Elston-Harris, 1992; Wharton & Marsh, 1978). 


albobalteata 




Psyttalia (=Opius) Common Indian egg-larval Braconid parasite, in Hawaii restricted to Melon 

fletcheri 

Fly.Also in the Philippines, Guam and int. to Ryukyu Is. (Clausen, 1956; 

Wharton & Gilstrap, 1983). From Thailand (Chinajariyawong, et al., 2000). 

Psyttalia (=Opius) A Braconid parasite from SE Asia (Wharton & Gilstrap, 1983). From Thailand 

makii 


Opius oophilus A Braconid parasite from SE Asia (Cochereau, 1970). 

Bactrocera tau Opius angaleti A Braconid larval-pupal parasite from N. Borneo (Narayanan & Chawla, 

(Cont.) 

1962). 

Utetes bianchii A Braconid parasite from Thailand (Chinajariyawong, et al., 2000) 

Bactrocera 

thailandica 


Bactrocera Dipterophagus daci A Dipterophagid (Strepsiptera) gregarious (2-6 per host) endoparasite from 

tenuifascia 

Bactrocera 

trivialis 



Bactrocera tryoni Opius tryoni 

An Australian Braconid larval parasite present in small numbers (Snowball, et 

(= Biosteres tryoni) al., 1962; May & Kleinschmidt, 1954), 6.2% (Allman, 1939).Has a wide host 

range (Clausen, 1956; Silvestri, 1914; Wharton & Gilstrap, 1983; Wharton & 

Marsh, 1978; Cochereau, 1970; Narayanan & Chawla, 1962) 

Opius fijiensis An Australian, S. Pacific rare Braconid egg-larval parasite (Snowball, et al., 

1962; Wharton & Gilstrap, 1983; Cochereau, 1970). 

Opius deeralensis An Australian Braconid egg-larval parasite present in small numbers 

(Snowball, et al., 1962; Wharton & Gilstrap, 1983; Cochereau, 1970). 


PPQ




Bactrocera 

tsuneonis 

Bactrocera 

tuberculata 

Bactrocera 

umbrosa 

Bactrocera 

verbascifoliae 

Opius sp. ? africanus A Braconid larval parasite from Australia (May & Kleinschmidt, 1954; 


Opius fletcheri A Braconid larval parasite from Australia (May & Kleinschmidt, 1954; 


Opius kraussi A Braconid larval parasite from Australia (May & Kleinschmidt, 1954; 


Fopius schlingeri A egg-pupal Braconid parasite from Australia (Wharton, 1999) 

Biosteres 

longicaudatus 

Biosteres 

vandenboschi 

An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, 

Australia,America, Carribean, Florida (Wharton & Gilstrap, 1983; Snowball, 

1966; Wharton & Marsh, 1978). 






(=Opius oophilus 

A Braconid egg/pupal parasite from SE Asia & dominant in Hawai Females 

also carry bacteria and fungi that destroy eggs(Bosch & Haramoto, 1951; 

Clausen, 1956; Snowball, 1966).Intr. to Australia (Cochereau, 1970; Quimio 

& Walter, 2001). 

Opius longicaudatus An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji, 

Australia,America, Carribean, Florida (Wharton & Gilstrap, 1983; Snowball, 

1966). 

var. taiensis From SE Asia, intr. to Australia (Cochereau, 1970). 




Opius froggatti A Braconid larval parasite in Australia, New Caledonia described from 

numerous specimens (Cochereau, 1970). 

Pachyneuron 

A Pteromalid parasite from SE Asia (Cochereau, 1970). 

vindemmiae 


longicaudata 






longicaudata 



Fopius vandenboschi A Braconid parasite from Malaysia and Thailand (Chinajariyawong, et al., 

2000). 

Phaenocarpa 

bactrocerae 

A Braconid parasite from the Phillippines (Thompson, 1943; Narayanan & 

Chawla, 1962;Dammerman, 1929). 



longicaudata 




PPQ


Bactrocera 

xanthodes 

Bactrocera 

zonata 

Carpomya 

incompleta 

Capparimyia 

savastani 

Carpomya 

schineri 

Carpomya 

vesuviana 






fletcheri 



Gilstrap, 1983; Wharton & Marsh, 1978; Narayanan & Chawla, 1962). 


1983; Thompson, 1943). 

Tetrastichus 

giffardianus 

A Eulophid parasite from Hawaii-Fiji (Thompson, 1943; Simmons, 1935). 

Opius hageni A Braconid parasite from Fiji (Cochereau, 1970). 

Spalangia cameroni A Pteromalid larval-pupal parasite (Narayanan & Chawla, 1962). 

Spalangia sp. A Pteromalid parasite (Narayanan & Chawla, 1962). 

Syntomosphyrum A Eulophid parasite intr. Fiji from New South Wales (Narayanan & Chawla, 

indicum 

1962). 

Germalus pacificus A Lygaeid egg predator (Narayanan & Chawla, 1962). 

Opius fletcheri A Braconid predator (Narayanan & Chawla, 1962). 

Diachasmimorpha An Indo-Australian Braconid larval parasite, now in Hawaii, Fiji,America, 

(=Biosteres) 


longicaudatus Kapoor, 1993). Rate of parasitism 10% (Ghani, 1972). In Thailand 


Fopius (=Biosteres) 

persulcatus (?) 

A Braconid parasite from India (Wharton & Gilstrap, 1983). 


Austroopius sp. A Braconid parasite from India (Thompson, 1943). 

Opius sp. A Braconid parasite from India (Narayanan & Chawla, 1962). 



fletcheri 


Trybliographia daci A Cynipid larval-pupal parasite from Pakistan (Ahmad, et al., 1975). Rate of 

parasitism 45% (Ghani, 1972). 


(Wharton & Gilstrap, 1983; Thompson, 1943). 

Eupelmus urozonus A Eupelmid from Italy (Thompson, 1943). 

Tetrastichus sp. A Eupelmid from Eritrea (Thompson, 1943) 

Opius sp. A Braconid parasite from Israel (Fry, 1987). 


(Wharton & Gilstrap, 1983; Soria & Yana, 1959). 

Opius scrutator A Braconid parasite from Southern Europe, Central Asia (Kandybina, 1977). 

Opius rhagoleticola A Braconid parasite from Southern Europe, Central Asia (Kandybina, 1977). 

Habrocytus sp. A Pteromalid parasite from Southern Europe, Central Asia (Kandybina, 1977). 

Opius bianchii A Braconid larval-pupal parasite from northern India. (Fullaway,1951; 

Wharton & Gilstrap, 1983; Narayanan & Chawla 1962). 


PPQ




Carpomya 

vesuviana 

(Cont.) 

Carpophthoromyi 

a tritea 



A Braconid egg-larval parasite from the Indo-Australian Region (Wharton & 

Gilstrap, 1983; Quimio & Walter, 2001; Chinajariyawong, et al., 2000). 

Biosteres carpomyiae A Braconid larval parasite from India that attacks when the larvae respire at 

the fruit surface (Wharton &Gilstrap, 1983; Thompson, 1943; Narayanan & 

Chawla, 1962; Singh, 1989) introduced to Iran (Farrar & Chow, 2000). 

Fopius (= Biosteres) 

vandenboschi 

A Braconid 1st stage larval parasite from SE Asia, est. in Hawaii (Wharton & 

Gilstrap, 1983; Kapoor, 1993). From the Indian desert (Singh, 1989). From 

Thailand (Chinajariyawong, et al., 2000). 

Opius incisi A Braconid egg-larval parasite from SE Asia, est. in Hawaii (Wharton & 


Opius makii A Braconid parasite from SE Asia (Wharton & Gilstrap, 1983). 

Microbracon fletcheri A Braconid larval parasite which pupates outside the fruit pulp, from India 

(Thompson, 1943; Narayanan & Chawla 1962; Singh, 1989). 

Microbracon lefroyi A Braconid parasite from India (Thompson, 1943). 

Omphale sp.= A Eulophid parasite from India (Narayanan & Chawla, 1962). 


longicaudata 



Psyttalia sp. nr makii A Braconid parasite fromThailand (Chinajariyawong, et al., 2000). 


Biosteres caudatus A Braconid parasite from Africa (Wharton & Gilstrap, 1983). 

Biosteres fullawayi A Braconid parasite from Africa and widely spread to Hawaii, Spain, Puerto 

Rico, Australia (Wnarton & Gilstrap, 1983). 

Opius inconsuetus A Braconid parasite from Nigeria, est in Hawaii (?) (Wharton & Gilstrap, 

1983). 

Ceratitis spp. Biosteres sp. A Braconid parasite from Tanganyika (Thompson, 1943). 

Dirhinus ehrhorni A Chalcid parasite from S. Nigeria (Thompson, 1943) 

Dirhinus giffardii A Chalcid parasite from S. Nigeria (Thompson, 1943). 

Eupelmus sp. An Eupelmid parasite from Tanganyika (Thompson, 1943). 

Hedylus desideratus A Braconid parasite from Nigeria (Thompson, 1943; Narayanan & Chawla, 

1962). 

Isurgus sp. An Ichneumonid parasite from Tanganyika (Thompson, 1943. 

Microbracon sp. A Braconid parasite from Tanganyika (Thompson, 1943). 

Opius sp. A Braconid parasite from Tanganyika (Thompson, 1943). 

Sphegigaster sp. A Pteromalid species from Tanganyika (Thompson, 1943). 

Hedylus clypeatus A Braconid larval parasite (Narayanan & Chawla, 1962). 


dubius 


Spalangia afra A Pteromalid parasite from East Africa (Narayanan & Chawla, 1962). 

Pheidole 

megacephala 

A cosmopolitan Formicid predator (Narayanan & Chawla, 1962). 


PPQ




Ceratitis anonae Biosteres caudatus A Braconid larval parasite from Nigeria, French Cameroons, Sierra Leone 

(Silvestri, 1914; Wharton & Gilstrap, 1983; Narayanan & Chawla, 1962; 

Steck, et al., 1986). 

Galesus silvestrii A Diapriid pupal parasite in small numbers from Nigeria. Also found on 

Dirhinus giffardii (Silvestri, 1914; Nixon, 1930). 

Ceratitis 

antistictica 

Dirhinus giffardii (= 

Biosteres giffardii) 

Ceratitis capitata Diachasmimorpha 

(=Opius) tryoni 

Ceratitis capitata 

(Cont.) 

A Braconid pupal parasite from Nigeria (Silvestri, 1914;(Wharton & Gilstrap, 

1983). 

Spalangia afra A Pteromalid pupal parasite from Nigeria (Silvestri, 1914; Boucek, 1963). 

Biosteres desideratus A Braconid parasite from Africa (Wharton & Gilstrap, 1983; Steck, at a, 

1986). 


Rico, Australia (Wharton & Gilstrap, 1983; Steck, et al., 1986). 



Tetrastichus giffardii A Eulophid egg-larval parasite from Dahomey, Nigeria, Victoria (Silvestri, 

1914). 

Biosteres caudatus A Braconid larval parasite from Nigeria (Silvestri, 1914; Wharton & Gilstrap, 

1983). 


kraussii 

Opius fullawayi 

(=Biosteres fullawayi) 

An Australian Braconid egg-larval parasite from Queensland fruit fly now 

dominant on Medfly in Hawaii (Clausen, 1956; Adams, 1994; Wharton & 

Gilstrap, 1983; Thompson, 1943; Wharrton & Marsh, 1978; Cochereau, 

1970; Fry, 1987;Narayanan & Chawla, 1962; Vargas, 2002; Duan & 

Messing, 2003; Ovruski, et al., 2000; Vargas et al., 2000). 

An Australian Braconid larval parasite. Unsuccessfully released in Hawaii 

against Medfly in 1947-52 (Rungrojwanich & Walter, 2000; Messing, et al., 

2000). 

A West African Braconid egg-larval parasite from Ceratitus spp, effective on 

Medfly in Hawaii (Clausen, 1956; Wharton & Gilstrap, 1983). A Braconid 

parasite from Africa and widely spread to Hawaii, Spain, Puerto Rico, 

Australia (Wharton & Gilstrap, 1983; Thompson, 1943). 

Opius hirtus A Braconid parasite from Costa Rica (Ovruski, et al., 2000) 

Opius humilis A South African Braconid larval parasite from Medfly effective on Medfly in 

Hawaii where it has since been displaced by Diachasmimorpha. Also in 

Bermuda (Clausen, 1956; Silvestri, 1914; Wharton & Gilstrap, 1983; 

Thompson, 1943; Narayanan & Chawla, 1962; Vargas, 2002). 

Dirhinus ehrhorni A Chalcid parasite (?) from Nigeria (Silvestri, 1914). 

Dirhinus giffardii A West African Chalcid pupal parasite, only occasionally found in Hawaii, also 

in Egypt & Brazil (Clausen, 1956; Silvestri, 1914; Thompson, 1943; 

Cochereau, 1970; Narayanan & Chawla, 1962). Intro. Reunion (Etienne, 

1973). 

Tetrastichus 

giffardianus 

A West African Eulophid pupal parasite, almost negligible, except in certain 

fruits. (Clausen, 1956; Thompson, 1943).Intro. Reunion, Hawaii (Etienne, 

1973; Cochereau, 1970; Narayanan & Chawla, 1962). 

Opius perproximus A West African Braconid larval parasite bred in lab (Silvestri, 1914; Wharton 

& Gilstrap, 1983). 

Galesus silvestrii A Diapriid pupal parasite from Nigeria bred in lab (Silvestri, 1914; Thompson, 

1943). 


PPQ





(Cont.) 

Trichopria capensis A Diapriid pupal parasite from South Africa (Silvestri, 1914). 


indicum 

An Eulophid late instar larval parasite from India of strong potential. Also 

said to parasitize Bactocera & Dacus (Silvestri, 1914; Thompson, 1943). 

Intro. Reunion (Etienne, 1973). 

Opius kraussii A Braconid larval parasite from Australia with decided preference for Medfly 

(Fullaway, 1951). 


(=Biosteres arisanus ) 


A Braconid egg/pupal parasite which outcompetes other parasites (Wood, 

1998; Wharton & Gilstrap, 1983; Fry, 1987) in Costa Rica (Ovruski, et al., 

2000). 

A Braconid egg/pupal parasite from SE Asia intr. to Hawaii (Cochereau, 

1970). 

The above are synonyms ( Quimio & Walter, 2001). 

Bracon celer A Braconid larval parasite from Kenya (Wharton, et al., 2000). 

Doryctobracon 

areolatus 


in Florida (Wharton & Gilstrap, 1983; Leonel, et al., 1995; Katiyar, et al., 

1995) in Argentina, Venezula, Brasil (Ovruski, et al., 2000) 

Coptera haywardi A Diapriid pupal endoparasitoid from Latin America, but unable to develop on 

irradiated hosts (Menezes, et al., 1998; Sivinski, et al., 1998). 

Coptera occidentalis A Diapriid pupal endoparasitoid from California, exported to Slovakia and 

reared in lab on Medfly (Kazimirova & Vallo, 1999) 

Coptera robustior A Diapriid pupal endoparasitoid from Kenya (Wharton, et al., 2000; as 

Coptera sp.). Also in Guinea, Nigeria, South Africa (Yoder & Wharton, 

2002). 



Aganaspis pelleranoi An Eucoilid parasite from Costa Rica & Argentina which seeks out late instar 

larvae through fruit holes (Sivinski, et al., 1997; Ovruski, 1994; Ovruski, et 

al., 2000; Ovruski, 2002) 

Aganaspis 

A parasite from Costa Rica (Ovruski, et al., 2000) 

nordlanderi 

Biosteres desideratus A Braconid parasite from Africa, reared in lab (Wharton & Gilstrap, 1983). 

Odontosema 

anastrephae 

Lopheucoila 

anastrepha 


indica 

Trichopria 

anastrephae 

A non-specific Cynipid larval-pupal parasite from Costa Rica (Wharton, et al., 

1981; Ovruski, et al., 2000) 


(Gallardo & Diaz, 1999; Ovruski, et al., 2000)) 

A parasite from Argentina (Ovruski, et al., 2000). Mexico, Peru, Bolivia (Rogg 

& Camacho, 2003). 

A parasite from Argentina (Ovruski, et al., 2000) 

Utetes anastrephae A parasite from Argentina (Ovruski, et al., 2000) 

Doryctobracon 

areolatus 

A Braconid larval-pupal parasite in Argentina (Ovruski, 2002) In Brazil 

(Marchiori, et al., 2000). 

Chiracanthium mildei A sac spider which may be an important nocturnal predator of Medfly in 

Israel, as it is strongly attracted to the odor of medfly males (Kaspi, 2000). 

Ganaspis carvalhoi A non-specific Cynipid larval-pupal parasite from Costa Rica (Wharton, et al., 

1981) 


PPQ



(Cont.) 



Biosteres 


(= Diachasmimorpha) 

longicaudatus 

1981; Wharton & Gilstrap, 1983; Wharton & Marsh, 1978; Narayanan & 

Chawla, 1962). Intro. Reunion (Etienne, 1973; Fry, 1987) in Guatemala 

(Ovruski, et al., 2000) In Bolivia (Rogg & Camacho, 2003). 

Biosteres oophilus A Braconid egg-pupal parasite introduced into Costa Rica (Wharton, et al., 

1981; Wharton & Marsh, 1978). 

Fopius ceratitivorus A larval-pupal Braconid parasite from Africa (Wharton, 1999); from Kenya 

(Wharton, et al., 2000). 

Fopius caudatus A larval-pupal Braconid parasite From Kenya (Wharton, et al., 2000). 

Biosteres angaleti A Braconid egg parasite from Indo-Australia (Wharton & Gilstrap, 1983). 

Biosteres bevisi A Braconid parasite from the South African Region reared from Medfly in 

Hawaii (Wharton & Gilstrap, 1983). 

Biosteres dacusii A Braconid parasite from India, reared from Medfly in Hawaii (Wharton & 

Gilstrap, 1983; Cochereau, 1970; Narayanan & Chawla, 1962). 


Rico, Australia (Wnarton & Gilstrap,1983). 

Biosteres hageni A Braconid parasite from Fiji reared in lab (Wharton & Gilstrap, 1983). 

Biosteres 

vandenboschi 



Opius bellus A Braconid parasite from S. and C. America, also Morocco where 28% 

parasitism exists (Wharton & Gilstrap, 1983; Leonel, et al., 1995; 

Cochereau, 1970; Narayanan & Chawla, 1962; Ovruski, et al., 2000)). 

Psyttalia (=Opius) 

concolor 

A Braconid parasite from northern Africa, intr. throughout the Mediterranean 

(Wharton & Gilstrap, 1983; Narayanan & Chawla, 1962; Kimani-Njogu, et al., 

2000). Intro. Reunion (Etienne, 1973), in Costa Rica (Ovruski, et al., 2000) 

Opius perproximus A Braconid larval parasite from West Africa, in lab (Wharton & Gilstrap, 

1983). 

Pachycrepoideus A Pteromalid parasite in Hawaii (Thompson, 1943). 

dubius 


(= Pachyneuron) 

vindemmiae 


(Etienne, 1973; Narayanan & Chawla, 1962). In Argentina (Ovruski, 2002). 

In Bolivia (Rogg & Camacho, 2003). 

Pimpla pomorum An Ichneumonid parasite from Spain (Thompson, 1943). 

Psyttalia distinguenda A Braconid pupal parasite from Reunion, Mauritius, Madagascar (Wharton, 

et al., 1999). 

Psyttalia 

A Braconid pupal parasite from Reunion Island (Wharton, et al., 1999). 

insignipennis 

Tetrastichus sp. An Eulophid parasite from Hawaii (Thompson, 1943). 

Spalangia endius A Chalcid larval-pupal (?) parasite with many hosts, world-dist. (Boucek, 

1963). 

Iridomyrmex humilis Predation by the Argentine ant (Formicidae) from 2.3, 38.5 to 50% of last 

stage larvae, pupae and ternal adults in Hawaii (Wong, et al., 1984). 

Solenopsis geminata Predation by the fire ant (Formicidae) in Guatemala on falling larvae and 

pupae from 7-25% (Eskafi & Kolbe, 1990); in Hawaii, 73.4% (Wong & Wong, 

1988). 

Opius longicaudatus A Braconid larval parasite from SE Asia intr. to Hawaii (Cochereau, 1970). 

var. malaiensis 


PPQ




Eucoila sp. A Cynipid parasite from Mauritius (Orian & Moutia, 1960). 

Isurgus sp. An Ichneumonid parasite from Africa (Narayanan & Chawla, 1962). 

Aphaereta minuta A Braconid larval-pupal parasite (Narayanan & Chawla, 1962). 

Opius fletcheri A Braconid larval-pupal parasite (Narayanan & Chawla, 1962). 

Trybliographa sp. A Cynipid entoparasitoid from Malaya (Narayanan & Chawla, 1962). 

Trybliographa daci A Cynipid endoparasitoid from Pakistan, also parasitizing Dacus dorsalis 

(Cals-Usciati, et al., 1985) 

Spalangia 


A Pteromalid pupal parasite est. in Hawaii (Narayanan & Chawla, 1962). 

Sphegigaster sp. A Pteromalid parasite (Narayanan & Chawla, 1962). 

Belonuchus 

A Staphylinid larval predator from Brazil, reared ex lab (in bitter orange) in 

rufipennis 

Italy (Narayanan & Chawla, 1962). 

Doryctobracon A Braconid parasite from Venezuela (Katiyar, et al., 1995) from Guatemala 

crawfordi 

(Ovruski, et al., 2000) 

Opius incisi A Braconid larval parasite found ex Medfly in Hawaii (Clancy, 1952). 

Pheidole 

An ant (Formicidae) predator, which kills large numbers of larvae (Kapoor, 

megacephala 

1993). 

Ectemnius 

An adult Crabronine wasp predator which store the prey as food for the young 

timidoventris 

(Kapoor, 1993). 

Crabo unicolor An adult Crabronine wasp predator which store the prey as food for the young 

(Kapoor, 1993). 

Ceratitis catoirii Psyttalia 

A Braconid parasite from Mauritius which apparently controls this pest (Orian 

insignipennis 

& Moutia, 1960; Wharton, et al., 1999). 

Ceratitis colae Galesus silvestrii A Diapriid pupal parasite from Nigeria bred in lab (Silvestri, 1914) 

Splangia afra A Pteromalid pupal parasite from Nigeria bred in lab (Silvestri, 1914; Boucek, 

1963). 

Tetrastichus giffardii A widely diffused Chalcid egg-larval parasite from Nigeria, Dahomey, Victoria, 

Tanganyika (Silvestri, 1914; Thompson, 1943). Introduced to Mexico, to 

Peru to Bolivia (Rogg & Camacho, 2003). 

Aeromyrma vorax An ant (Formicidae) predator of pupae (Silvestri, 1914). 

Opius perproximus A Braconid larval parasite from West Africa (Wharton & Gilstrap, 1983). 



Ceratitis 

contramedia 

Biosteres giffardii A Braconid parasite from Africa (Wharton & Gilstrap, 1983). 



Ceratitis cosyra Opius cosyrae A Braconid parasite from northern Africa, intr. through the Mediterranean 

(Wharton & Gilstrap, 1983; Thompson, 1943; Narayanan & Chawla, 1962). 

Opius perproximus A Braconid larval parasite from West Africa (Silvestri, 1914; Wharton & 

Gilstrap, 1983; Narayanan & Chawla, 1962; Steck, et al., 1986). 

Biosteres fullawayi A Braconid parasite from the Cameroons (Steck, et al., 1986). 

Ceratitis cosyra Coptera silvestrii A Psiline parasite from Africa (Benin, Ghana, Niger, Nigeria, Senegal, Kenya, 

(Cont.) 



PPQ




Ceratitis 

ditissima 

Biosteres caudatus A Braconid larval parasite from Africa (Wharton & Gilstrap, 1983). 

Biosteres giffardii A Braconid parasite from Africa (Wharton & Gilstrap, 1983). 

Ceratitis 

flexuosa 

Opius perproximus A Braconid parasite from the Cameroons (Steck, et al., 1986). 

Ceratitis griffardi Dirhinus erhorni A possible Chalcid pupal parasite from Nigeria (Silvestri, 1914). 

Splangia afra A Pteromalid pupal parasite bred in lab (Silvestri, 1914; Boucek, 1963). 

Opius perproximus A Braconid larval parasite from West Africa (Silvestri, 1914; Wharton & 


Diachasma 

A Braconid pupal parasite from French Guinea (Silvestri, 1914).1983).A 

fullawayi(=Biosteres Braconid parasite from Africa and widely spread to Hawaii, Spain, Puerto 

fullawayi) 


Hedylus 

A Braconid larval parasite from French Guinea (Silvestri, 1914; Wharton & 

giffardii(=Biosteres 

giffardii) 


Opius perproximus 

var. modestior 

A Braconid parasite from Nigeria (Silvestri, 1914). 

Biosteres caudatus A widely diffused Braconid larval parasite from Victoria (Silvestri, 1914; 

Wharton & Gilstrap, 1983; Narayanan & Chawla, 1962). 

Tetrastichus giffardii A widely diffused Eulophid egg-larval parasite from Nigeria, Dahomey, Victoria 

(Silvestri,1914; Thompson, 1943). 

Ceratitis 

malgassa 

Galesus silvestrii A Diapriid pupal parasite from Nigeria (Silvestri, 1914; Nixon, 1930). 

Ceratitis 

pedestris 


Ceratitis 

Coptera robustior = A Psiline pupal parasite from Kenya, Guinea, Nigeria, South Africa (Yoder & 

punctata (Galesus silvestrii var. Wharton, 2002). A Diapriid pupal parasite from French Guinea (Silvestri, 

robustior) 

1914; Nixon, 1930). 

Hedylus giffardii(= A French Guinean Braconid larval (?) parasite (Silvestri, 1914; (Wharton & 

Biosteres giffardii) Gilstrap, 1983; Narayanan & Chawla, 1962). 

Diachasma fullawayi A Braconid pupal parasite from French Guinea (Silvestri, 1914; Wharton & 

var. robustum = Gilstrap.1983). A Braconid parasite from Africa and widely spread to Hawaii, 

Biosteres fullawayi) Spain, Puerto Rico, Australia (Wharton & Gilstrap, 1983). 

Dorylus (Anomma) 

nigricans var. hybrida 

A Formicid predator of pupae from West Africa (Silvestri, 1914). 

Ceratitis quinaria 

Opius perproximus A Braconid larval parasite from West Africa, Cameroons. (Wharton & Gilstrap, 

1983; Steck, et al., 1986). 

Ceratitis rosa Fopius ceratitivorus A larval-pupal Braconid parasite from Africa (Wharton, 1999) 

Opius africanus A Braconid larval parasite from South Africa (Silvestri, 1914; Wharton & 


Pachyneuron 

A Pteromalid pupal parasite from Morocco (Thompson, 1943).Intro. Reunion 

vindemmiae 



in Egypt & Brazil (Clausen, 1956; Silvestri, 1914; Thompson, 1943). Intro. 

Reunion (Etienne, 1973). 


PPQ





Ceratitis rosa 

(Cont.) 


Ceratitis rubivora 



Ceratitis simi Coptera silvestrii A Psiline parasite from Africa (Benin, Ghana, Niger, Nigeria, Senegal, Kenya, 


Ceratitis stictica Tetrastichus giffardii An Eulophid parasite from W. Africa (Thompson, 1943). 

Ceratitis tritea Tetrastichus oxyurus A Eulophid pupal (?) parasite from Nigeria (Silvestri, 1914). 

Opius inconsuetus A Braconid parasite from Nigeria (Silvestri, 1914). 

Diachasma fullawayi A pupal parasite from Nigeria (Silvestri, 1914). 

Biosteres caudatus A larval parasite from Nigeria (Silvestri, 1914). 

Dacus spp. Biosteres desideratus A Braconid parasite from Africa (Wharton & Gilstrap, 1983; Bridwell, 1918; 

Thompson, 1943; Narayanan & Chawla, 1962). 

Biosteres 

ottotomoanus 

A Braconid parasite from Cameroon (Wharton & Gilstrap, 1983). 

Bathyaulax sp. A Braconid parasite from India (Thompson, 1943). 

Biosteres formosanus A Braconid parasite from Formosa (Thompson,1943). 

Tetrastichus 

A Eulophid parasite from the S. Pacific & Australia (Thompson, 1943; 

giffardianus 


Dirhinus auratus A Chacid pupal parasite from Pakistan, SE Asia, Mauritius (Narayanan & 


Spalangia cameroni A Chalcid, pro larval-pupal parasite from Europe, Africa, Asia,& S. America, 

pro imported to Pacific Is.Many diverse hosts (Boucek, 1963; (Narayanan & 


Opius fletcheri A Braconid parasite (Narayanan & Chawla, 1962). 

Opius skinneri A Braconid larval-pupal parasite from the Phillippines (Narayanan & Chawla, 

1962). 

Opius tryoni A Braconid parasite (Narayanan & Chawla, 1962). 

Opius watersi A Braconid parasite (Narayanan & Chawla, 1962). 

Phaenocarpa leveri A Braconid parasite from Fiji (Narayanan & Chawla, 1962). 

Opius angaleti A Braconid from North Borneo (Narayanan & Chawla, 1962). 

Dirhinus giffardii A Chalcid parasite intr. to Fiji (Narayanan & Chawla, 1962). 

Trichopria capensis A Diapriid parasite (Narayanan & Chawla, 1962). 

Opius fullawayi An Ichneumonid parasite from Africa (Narayanan & Chawla, 1962). 

Opius perproximus A Braconid parasite (Narayanan & Chawla, 1962). 

Dacus axanus 



Dacus bellulus Dipterophagus daci A Dipterophagid (Strepsiptera) gregarious (2-6 per host) endoparasite from 

Australia which attacks the soft ternal adult stage, resulting in sterile hosts 



PPQ


Dacus bivittatus Diachasma fullawayi 

var. robustum 



A pupal parasite from French Guinea (Silvestri, 1914). 

Biosteres caudatus A widely diffused Braconid larval parasite from Nigeria (Silvestri, 1914; 

Steck, et al., 1986). 

Coptera silvestrii 

(=Galesus silvestrii) 

Tetrastichus 

giffardii(=Biosteres 

giffardii) 

A Diapriid pupal parasite from Nigeria bred in lab (Silvestri, 1914). A Psiline 

parasite from Africa (Benin, Ghana, Niger, Nigeria, Senegal, Kenya, 


A widely diffused Braconid egg-larval parasite from Victoria (Silvestri, 1914; 

Wharton & Gilstrap, 1983; Thompson, 1943). 

Tetrastichus dacicida A Eulophid from the Cameroons (Thompson, 1943; Narayanan & Chawla, 

1962). 

Spalangia afra A Pteromalid (pupal?) parasite from Africa (Boucek, 1963; Narayanan & 


Dirhinus sp. A Chacid parasite from Africa (Narayanan & Chawla, 1962). 

Biosteres desideratus A Braconid parasite from the Cameroons (Steck, et al., 1986). 

Opius perproximus A Braconid parasite from the Cameroons (Steck, et al., 1986). 

Dacus ciliatus Opius perproximus A Braconid larval parasite from West Africa (Silvestri, 1914; Wharton & 

Gilstrap, 1983; Thompson, 1943; Narayanan & Chawla, 1962). 

Biosteres caudatus A widely diffused Braconid larval parasite from Dahomey (Silvestri, 1914; 


Biosteres giffardii An African Braconid larval parasite (Wharton & Gilstrap, 1983; Narayanan & 

Chawla, 1962; Ghani, 1972). 

Biosteres 

longicaudatus 


Caribbean, Florida (Wharton & Gilstrap, 1983; Wharton & Marsh, 1978). 



Opius fletcheri Common Indian Braconid egg-larval parasite, in Hawaii restricted to Melon 

Fly. Also in the Philippines, Guam and intro. to Ryukyu Is. (Clausen, 1956; 


Opius phaeostigma A Braconid parasite from South Africa (Wharton & Gilstrap, 1983; Thompson, 


Sarcophaga flagellata A Sarcophagid parasite (Diptera) from the Italian Somaliland (Thompson, 

1943). 

Diachasma brevistyli A Braconid parasite from the Italian Somaliland (Thompson, 1943). 

Spalangia afra A Chalcid (pupal?) parasite from Africa (Boucek, 1963; Narayanan & Chawla, 

1962). 

Opius brevistyli A Braconid parasite from Africa (Narayanan & Chawla, 1962). 

Opius incisi A Braconid parasite from India (Narayanan & Chawla, 1962). 

Dirhinus luzonensis The most abundant Chalcid parasite from India (Narayanan & Chawla, 1962). 

Pachycrepoideus A Pteromalid parasite (Narayanan & Chawla, 1962). 

dubius 

Spalangia grotiusi A Pteromalid parasite (Narayanan & Chawla, 1962). 

Spalangia 




PPQ




Dacus 

demmerezi 

Spalangia 


stomoxysiae 

Galesus sp. A Diapriid parasite from India (Narayanan & Chawla, 1962). 

Sarcophaga flagellata A dipterous Sarcophagid parasite (Narayanan & Chawla, 1962). 

Opius compensans A Braconid larval-pupal parasite from India which parasitizes 10-12% of the 

host (Narayanan & Chawla, 1962). 

Opius brevisi A braconid larval-pupal parasite from S. Africa which parasitizes 4-22% of the 

host (Daiber, 1966). 

Opius phaeostigma A Braconid parasite from South Africa (Wharton & Gilstrap, 1983; Thompson, 





Pachyneuron 

vindemmiae 

Biosteres 

longicaudatus 

Tetrastichus 

giffardianus 

Dacus frontalis Biosteres 

longicaudatus 




1981; Wharton & Gilstrap, 1983; Wharton & Marsh, 1978). Intro. Reunion 



fruits. (Clausen, 1956; Thompson, 1943). Intro. Reunion (Etienne, 1973). 

A Braconid parasite from the Cape Verde Islands (Fry, 1987). 

Opius concolor A Braconid parasite from the Cape Verde Islands (Fry, 1987). 

Dirhinus ?giffardii A Chalcid parasite from the Cape Verde Islands (Fry, 1987). 

Dacus humeralis Biosteres niger A Braconid parasite from Tanzania (Wharton & Gilstrap, 1983). 

Opius caudatus A Braconid parasite from the French Cameroons (Narayanan & Chawla, 

1962). 

Opius sp. A Braconid parasite (Narayanan & Chawla, 1962). 

Tetrastichus giffardii An Eulophid parasite from Tanganyika (Narayanan & Chawla, 1962). 

Dacus 

longicornis 


Dacus 

longistylus 

Dorylus affinis An ant (Formicidae) predator of fruit fly pupae from Africa (Silvestri, 1914) 

Opius dexter A Braconid parasite from Senegal (Silvestri, 1914; Wharton & Gilstrap, 

1983). 

Dacus 

lounsburyii 

Austroopius sp. A Braconid parasite from India (Thompson, 1943). 

Dacus 

momordicae 

Biosteres caudatus A Braconid parasite from Africa (Wharton & Gilstrap, 1983). 

Dacus 

Tetrastichus 

An Eulophid parasite (Narayanan & Chawla, 1962). 

punctatifrons giffardianus 

Dacus 

Opius angaleti A Braconid larval-pupal parasite from North Borneo (Narayanan & Chawla, 

smieroides 

Dacus 

solomonensis 

1962). 


PPQ



Species 

Dacus telfaireae 

Dacus 

vertebratus 

Dirioxa pornia 

Parasites/Predators Notes 

Dirioxa confusa 

Epochra 

canadensis 

Opius oophilus A Braconid egg parasite from SE Asia & dominant in Hawaii. Females also 

carry bacteria and fungi that destroy eggs (Bosch & Haramoto, 1951; 

Clausen, 1956; Snowball, 1966). 

Euphranta 

connexa 

Opius testaceus A Braconid parasite from Europe, Russia (Kandybina, 1977). 

Euphranta 

turpiniae 


Eurosta 

solidaginis 

Mordellistena sp. A Coleopterous (Mordellidae) predator (parasite??) from the USA (Fry, 1987). 

Eurytoma sp. A Eurytomid parasite from the USA (Fry, 1987). 

Eutreta 


xanthochaeta 

Gilstrap, 1983; Wharton & Marsh, 1978.). 

Gonioglossum Opius testaceus A Bracinid parasite from Central, Southern Russia, Europe (Kandybina, 

wiedemanni 

1977). 

Lonchaea 

pendus 

Monacrosthcius 

citricola 

Myiopardalis 

pardalina 

Eucolia weldi A Cynipid parasite from Brazil (Narayanan & Chawla, 1962). 

Myoleja limata Biosteres melleus A Braconid parasite from Minnesota and Nova Scotia to Florida (Wharton & 

Marsh, 1978). 

Myoleja lucida Opius rhagoleticola A Braconid parasite from Russia, Switzerland (Kandybina, 1977; Fry, 1987). 

Halticoptera laevigata A Pteromalid parasite from Russia, Germany, Switzerland (Kandybina, 1977; 

Fry, 1987; Gienapp, 2000). 

Habrocytus sp. A Pteromalid parasite from Siberia (Kandybina, 1977). 

Phygadeuon elegans An Ichneumonid parasite from Switzerland (Fry, 1987). 

Phygadeuon 

wiesmanni 

An Ichneumonid parasite from Austria, Germany, Switzerland (Fry, 1987). 

Phygadeuon sp. An Ichneumonid parasite from Siberia (Kandybina, 1977). 

Gelis sp. An Ichneumonid parasite from Siberia, Switzerland (Kandybina, 1977; Fry, 

1987). 

Myoleja sinensis Opius myolejae A Braconid parasite from Russia (Kandybina, 1977). 

Halticoptera sp. A Pteromalid parasite from Russia (Kandybina, 1977). 

Nitrariomyia Habrocytus sp. A Pteromalid pupal parasite from arid zones of Central Asia (Kandybina, 

lukjanovitshi 

1977). 

Pardalaspis Spalangia gemina A Chalcid pupal parasite with a broad host range from Asia through S. 

cyanescens 

America (Boucek, 1963). 

Procecidochares Biosteres tryoni A Braconid egg-larval parasite from eastern Australia, est. Hawaii (Wharton & 

utilis 



PPQ




Rhagoletis sp. Salticus scenicus An Aranead predator from British Columbia (Fry, 1987). 

Rhagoletis 

alternata 

Opius rosae A Braconid parasite (Kandybina, 1977). 

Opius rhagoleticola A Braconid parasite (Kandybina, 1977). 

Opius ophthalmicus A Braconid parasite form Far Eastern Russia (Kandybina, 1977). 

Opius alternatae A Braconid parasite from Far Eastern Russia (Kandybina, 1977). 

Opius sp. A Braconid parasite from Swizerland (Fry, 1987). 

Halticoptera sp. A Pteromalid parasite (Kandybina, 1977). 

Phygadeuon exiguus An Ichneumonid parasite from Germany, Switzerland (Fry, 1987). 

Phygadenon sp. An Ichneumonid parasite from Western Siberia (Kandybina, 1977). 

Gelis sp. An Ichneumonid parasite from Western Siberia (Kandybina, 1977). 

Rhagoletis 

basiola 

Halticoptera rosae An Pteromalid egg-larval parasitoid from Europe (Hoffmeister, 2000) 

Rhagoletis 

batava 


Thyridanthrax after A Bombyliid parasite (Kandybina, 1977). 

Habrocytus sp. A Pteromalid parasite (Kandybina, 1977). 

Phygadenon sp. An Ichneumonid parasite (Kandybina, 1977). 

Gelis sp. An Ichneumonid parasite (Kandybina, 1977). 

Rhagoletis 

berberidis 

Habrocytus sp. A Chalcid parasite from Europe, Russia (Kandybina, 1977). 

Eurytoma sp. A Chalcid parasite from Europe, Russia (Kandybina, 1977). 

Eupelmus sp. A Chalcid parasite from Europe, Russia (Kandybina, 1977). 

Opius rhagoleticola A Braconid parasite from Europe, Russia - Rare (Kandybina, 1977). 

Rhagoletis 

boycei 

Biosteres juglandis A Braconid parasite from Arizonia, New Mexico (Wharton & Marsh, 1978). 

Rhagoletis 

caucasica 

Opius scrutator (?) A probable Braconid parasite (Kandybina, 1977). 

Rhagoletis Opius rhagoleticola A Braconid larval-pupal parasite from Siberia, Europe (Kandybina, 1977; 

cerasi 

Vogel, 1950; Fry, 1987;Narayanan & Chawla, 1962). 

Opius testaceus A Braconid parasite (Kandybina, 1977). 

Opius sayanicus A Braconid parasite (Kandybina, 1977). 

Opius magnus A Braconid full-grown larval parasite (10-30% on wild cherry only) from Italy 

(Fry, 1987; Monaco, 1984). 

Opius sp. A Braconid parasite from Switzerland (Fry, 1987). 

Cremnodes 

articapillus 

An Ichneumonid parasite from Austria, Switzerland (Fry, 1987). 

Habrocytus sp. A Pteromalid pupal parasite from Siberia (Kandybina, 1977) 

Coptera occidentalis A Diapriid parasite from the Pacific Coast of the USA which overwinters in 

diapausing R. cerasi (Kazimirova & Vallo, 1999) 

Halticoptera laevigata A Pteromalid parasite from Switzerland (Fry, 1987). 

Halticoptera sp. A Pteromalid parasite (Kandybina, 1977). 

Phygadeuon elegans An Ichneuomid parasite from Switzerland (Fry, 1987). 


PPQ


Rhagoletis 

cingulata 

Rhagoletis 

completa 

Rhagoletis 

completa(Cont.) 

Rhagoletis 

conversa 

Rhagoletis 

fausta 

Phygadeuon 

scaposus 

Phygadeuon 

wiesmanni 



An Ichneumonid parasite from Czechoslovakia (Fry, 1987). 

An Ichneumonid parasite from Austria, Germany, Switzerland which may 

attack larvae when entering ground to pupate; locally dominant @ 72.5% of 

pupae (Vogel, 1950; Fry, 1987; Narayanan & Chawla, 1962; Boller, 1966). 

Phygadeuon sp. An Ichneumonid pupal parasite from Siberia (Kandybina, 1977). 

Gelis sp. Ichneumonid pupal parasites from Siberia, Austria, Switzerland (Kandybina, 

1977; Fry, 1987). 

Gelis bremeri An Ichneumonid pupal parasite from Europe (Vogel, 1950; Narayanan & 


Polypeza forsteri A Diapriid parasite from Europe (Vogel, 1950;Narayanan & Chawla, 1962; 

Boller, 1966). 

Belonuchus 

rufipennis 

A Staphylinid larval predator from Brazil reared in lab (in cherries) in Italy 


Myrmica laevinodis A locally dominant ant (Formicidae) predator in Switzerland, with larval losses 

of 35-63% when migrating to the soil (Boller, 1966). 

Anisodactylus 

binotatus 

A local carabid predator in Switzerland with larval and pupal predation (Boller, 

1966). 

Paedrus litoralis A local staphid predator in Switzerland with larval and pupal predation (Boller, 

1996). 

Coptera cingulatae A Diapriid parasite from eastern USA, west to Kansas (Muesebeck, 1980). 

Coptera occidentalis A Diapriid parasite from the Pacific Coast of the USA (Muesebeck, 1980). 

Biosteres melleus A Braconid parasite from Minnesota and Nova Scotia to Florida (Wharton & 

Marsh, 1978). 

Diachasma (Opius) 

ferrugineum 

A Braconid parasite fromNorth America (Wharton & Marsh, 1978; Narayanan 

& Chawla, 1962). 



Opius humilus A Braconid larval parasite from South Africa, est. in Hawaii, Bermuda 


Coptera evansi A Diapriid parasite from SW USA (Muesebeck, 1980). 

Coptera occidentalis A Diapriid parasite from the Pacific Coast of the USA (Muesebeck, 1980). 

Biosteres sublaevis A Braconid parasite from Texas (Wharton & Marsh, 1978). 

Tetrastichus 

giffardianus 

A Eulophid parasite (Narayanan & Chawla, 1962). 



Diachasma 

ferrugineum 

A Braconid parasite fromNorth America (Wharton & Marsh, 1978). 

Eucoila sp. A Cynipid larval-pupal parasite (Narayanan & Chawla, 1962). 


PPQ




Rhagoletis 

ferruginea 

Rhagoletis 

flavicincta 

Rhagoletis 

indifferens 

Rhagoletis 

juglandis 

Rhagoletis 

kurentsovi 

Rhagoletis 

lycopersella 

Rhagoletis 

magniterebra 

Rhagoletis 

meigeni 

Rhagoletis 

mendax 

Rhagoletis nova 

Rhagoletis 

pastranai 

Rhagoletis 

pomonella 


dubius 

A Pteromalid larval-pupal parasite (Narayanan & Chawla, 1962). 

Tetrastichus faustus A Eulophid parasite (Narayanan & Chawla, 1962). 


vindemiae 

A Pteromalid generalist ectoparasitoid pupal parasite from Africa and India, 

introduced into Hawaii, Mexico and Central America (Guillen, et al., 2002). 

Opius bellus A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995; Ovruski, 

et al., 2000). 


(?) 

Opius scrutator A Braconid parasite (Kandybina, 1977). 

Diachasma muliebre A Braconid parasite from Washington to California (Wharton & Marsh, 1978; 



vindemiae 

A Pteromalid pupal parasite from Washington with a 20-37% parasitism rate 

(Burditt & White, 1987) A Pteromalid generalist ectoparasitoid pupal parasite 

from Africa and India, introduced into Hawaii, Mexico and Central America 

(Guillen, et al., 2002). 


Biosteres juglandis A Braconid parasite from Arizona, New Mexico (Wharton & Marsh, 1978). 


juglandis 

A Braconid larval parasite from Arizona (Henneman, et al., 2002) 

Opius kurentzov A Braconid parasite from Russia (Kandybina, 1977). 

Halticoptera sp. A Pteromalid parasite from Russia (Kandybina, 1977). 


Phygadeuon 

wiesmanni 

Biosteres (Opius) 

melleus 

An Ichneumonid parasite from Austria (Fry, 1987). 

A Braconid parasite from Minnesota and Nova Scotia to Florida. Said to be 

most effective parasite (Wharton & Marsh, 1978; Geddes, et al., 1987). 

Opius richmondi A Braconid parasite from Eastern Canada (Geddes, et al., 1987). 

Opius lectus A Braconid parasite from Eastern Canada (Geddes, et al., 1987). 

Doryctobracon A parasite from Brasil (Ovruski, et al., 2000) 

areolatus 

Doryctobracon A parasite from Brasil (Ovruski, et al., 2000) 

brasiliensis 

Opius bellus A parasite from Brasil (Ovruski, et al., 2000) 

Opius melleus A Braconid parasite (Clausen, 1956;Narayanan & Chawla, 1962; Monteith, 

1977). 

Biosteres melleus A Braconid larval-pupal parasite from Minnesota and Nova Scotia to Florida 

(Baranowski, et al., 1993; Wharton & Marsh, 1978; Gut & Brunner, 1994). 



PPQ


Rhagoletis 

reducta 

Rhagoletis 

ribicola 

Rhagoletis 

striatella 

Rhagoletis 

suavis 

Rhagoletis 

tabellaria 

Rhagoletis 

tomatis 

Rhagoletis 

turpiniae 



Coptera pomonellae A Diapriid parasite from eastern USA, west to Kansas and Minnesota 

(Muesebeck, 1980). 

Opius (Diachasma) 

alloeum = alloeus 

Diachasma 

ferrugineum = Opius 

ferrugineus 

A Braconid parasite fromNorth America. Parasitism is 30% with O. 

canaliculatus (Wharton & Marsh, 1978; AliNiazee, 1985;Narayanan & 

Chawla, 1962; Monteith, 1977). 

A Braconid parasite fromNorth America (Wharton & Marsh, 1978;Narayanan 

& Chawla, 1962; Monteith, 1977). 

Opius downesi A larval-pupal endoparasite from Washington ex 5% of pupae (AliNiazee, 

1985; Gut & Brunner, 1994; Narayanan & Chawla, 1962). 

Pteromalus sp. A larval ectoparasitoid from Washington (Gut & Brunner, 1994). 

Tetrastichus sp. Several larval ectoparasitoid species from Washington (Gut & Brunner, 

1994). 

Opius lectoides A Braconid larval-pupal endoparasite from Washington ex 0.5% of pupae 

(AliNiazee, 1985;Narayanan & Chawla, 1962). 

Opius canaliculatus ABraconid parasite from Quebec & New York which, with O. alloeum, has 30% 

early larval parasitism (AliNiazee, 1985). 

Aphaereta muscae A Braconid parasite (Kandybina, 1977; Narayanan & Chawla, 1962). 

Galesus sp. A Diapriid parasite (Kandybina, 1977; Narayanan & Chawla, 1962). 

Opius lectus A Braconid parasite from North America (Narayanan & Chawla, 1962; 

Monteith, 1977). 

Opius richmondi A Braconid parasite in North America ( Narayanan & Chawla, 1962). 

Patasson conotracheli A Mymarid egg parasite from Connecticut (Narayanan & Chawla, 1962). 

Opius ussyruensus A Braconid parasite from Russia and China (Kandybina, 1977). 

Chorebus sp. A Braconid parasite from Russia and China (Kandybina, 1977). 

Halticoptera sp. A Pteromalid parasite from Russia and China (Kandybina, 1977). 


Coptera pomonellae A Diapriid parasite from the Pacific Coast of the USA (Muesebeck, 1980). 

Biosteres melleus A Braconid parasite from Minnesota and Nova Scotia to Florida (Wharton & 

Marsh, 1978). 

Aganaspis pelleranoi A parasite from Mexico (Ovruski, et al., 2000) 

Biosteres near 

sublaevis 

A parasite from Mexico (Ovruski, et al., 2000) 

Dicerataspis spp. A parasite from Mexico (Ovruski, et al., 2000) 

Opius hirtus A parasite from Mexico (Ovruski, et al., 2000) 


PPQ




Rhagoletis 

zephyria 

Rhagoletis 

zephyria (Cont.) 

Rhagoletotrypeta 

astranai 

Opius downesi A larval-pupal Braconid endoparasite from Washington ex 2% of pupae 

(AliNiazee, 1985). 

Opius lectoides A larval-pupal Braconid endoparasite from Washington ex 2% of pupae 

(AliNiazee, 1985). 

Doryctobracon A solitary entoparasitoid from Brazil (Leonel, et al., 1995). 

areolatus 

Doryctobracon A solitary entoparasitoid from Brazil (Leonel, et al., 1995). 

brasiliensis 

Opius bellus A solitary entoparasitoid from Brazil (Leonel, et al., 1995). 

Utetes anastrephae A solitary entoparasitoid from Brazil (Leonel, et al., 1995). 

Tephritidae sp. Opius inquirendus A Braconid larval parasite from Victoria in an unknown host (Silvestri, 1914). 

Opius longicaudatus A Braconid larval parasite from the Philippines, India, Thailand, New 

Caledonia, Marianas (Fullaway, 1951). 

Opius formosanus A Braconidlarval parasite from Formosa (Fullaway, 1951). 

Opius compensans? A Braconid larval parasite from India (Fullaway, 1951). 

Opius watersi A Braconid larval parasite from India (Fullaway, 1951). 

Opius persulcatus A Braconid larval parasite from India, Philippines, Thailand (Fullaway, 1951). 

Opius deeralensis A Braconid larval parasite from Australia (Fullaway, 1951). 

Opius skinneri A Braconid larval parasite from the Philippines (Fullaway, 1951). 

Opius carpomyiae A Braconid larval parasite from India (Fullaway, 1951). 

Opius arisanus A Braconid larval parasite from India, Malaya, Thailand, Formosa (Fullaway, 

1951). 

Opius javanus A Braconid larval parasite from Thailand, Malaya (Fullaway, 1951). 

Opius fletcheri A Braconid larval parasite from India, Java, Malaya, Philippines (Fullaway, 

1951). 

Opius incisi A Braconid larval parasite from India, Malaya (Fullaway, 1951). 

Opius fijiensis A Braconidlarval parasite from Australia, Fiji, New Caledonia (Fullaway, 

1951). 

Opius makii A Braconid larval parasite from Formosa (Fullaway, 1951). 

Opius perkinsi A Braconid larval parasite from Australia (Fullaway, 1951). 

Opius froggatti A larval parasite from Australia (Fullaway, 1951). 

Opius manii A Braconid larval parasite from India, Malaya, Thailand (Fullaway, 1951). 

Terellia ruficauda Pronotalia sp. An Eulophid parasite from Germany (Fry, 1987). 

Tomoplagia sp. Opius itatiayensis A solitary entoparasitoid from Brazil (Leonel, et al., 1995). 

Tomoplagia 

rudolphi 

Opius tomoplagiae A solitary Braconid entoparasitoid from Brazil (Leonel, et al., 1995). 

Toxotrypana Doryctobracon A host specific larval-pupal Braconid parasite from Costa Rica, Mexico 

curvicauda toxotrypanae 

(Wharton, et al., 1981; Aluja, et al., 1990), El Salvador (Ovruski, et al., 

2000), Venezuela ( Martinez, et al., 1999). 

Coptera haywardi A Diapriid pupal parasite from Latin America (Sivinski, et al., 1998). 

Dihrinus giffardiii A Chalcid parasite intro. Puerto Rico (Narayanan & Chawla, 1962). 

Trirhithrum 

coffeae 

Fopius caudatus A dominant Braconid parasite from Kenya (Wharton, et al., 2000) 


PPQ


Trirhithrum 

coffeae (Cont.) 

Trirhithromyia 

cyanescens 

Trirhithrum 

inscriptum 

Trirhithrum 

nigerrimum 



Fopius ceratitivorus A Braconid parasite from Africa and widely spread to Hawaii, Spain, Puerto 


Biosteres caudatus A larval-pupal Braconid parasite from Africa (Wharton, 1999) 

Biosteres fullawayi A Braconid larval parasite from Africa (Wharton & Gilstrap, 1983). 

Opius sp. nr. 

desideratus 

A Braconid larval-pupal parasite from Uganda, in 7 to 23% of larvae 

(Greathead, 1972). 

Opius cosyrae A Braconid larval pupal parasite from Uganda, in 2-7% of larvae (Greathead, 

1972). 

Psilus sp. A Pteromalid pupal parasite from Uganda (Greathead, 1972). 

Tetrastichus giffardii A Eulophid multiple pupal parasite with up to 12 parasites per pupa, from 

Uganda (Greathead, 1972). 

Syntomosphyrum sp. An Eulophid pupal parasite from Uganda (Greathead, 1972). 

An Aleocharine A Staphylinid predator from Uganda (Greathead, 1972). 

Opius sp. A Braconid parasite from the Cameroons (Steck, et al., 1986). 

Diachasmimorpha An Australian Braconid egg-larval parasite in Reunion (Quilici, et al., 2000). 

tryoni 





(Wharton & Gilstrap, 1983). Intro. Reunion (Etienne, 1973). 

Biosteres 

longicaudatus 

Pachyneuron 

vindemmiae 


1981; Wharton & Gilstrap, 1983; Wharton & Marsh, 1978). Intro. Reunion 




Psyttalia fletcheri Common Indian egg-larvalBraconid parasite in Reunion (Quilici, etal., 2000). 

Tetrastichus 

giffardianus 


indicum 


fruits. (Clausen, 1956; Thompson, 1943).Intro. Reunion (Etienne, 1973). 

An Eulophid late instar larval parasite from India of strong potential. Also said 

to parasitize Bactocera & Dacus (Silvestri, 1914; Thompson, 1943). Intro. 


Opius insignipennis A Braconid larval-pupal parasite from Madagascar, Reunion Island (Orian & 

Moutia, 1960; Wharton, et al., 1999). 

Fopius okekai A Braconid pupal parasite from Kenya (Kimani-Njogu & Wharton, 2002). 

Opius perproximus 

var. modestior 

A Braconid parasite from Nigeria (Silvestri, 1914). 

Biosteres caudatus A Braconid larval parasite from Victoria (Silvestri, 1914; Wharton & Gilstrap, 

1983). 

Galesus silvestrii A Diapriid pupal parasite from Nigeria (Silvestri, 1914; Nixon, 1930). 



Microbracon celer A Braconid parasite from Kenya (Narayanan & Chawla, 1962). 



PPQ


Conservation of Predators and Parasites 


Trirhithrum 

nigrum 



Fobius okekai A Braconid pupal parasite from Kenya (Kimani-Njobu & Wharton, 2002) 

Opius perproximus A Braconid larval parasite from West Africa (Wharton & Gilstrap, 1983) 

Trirhithrum 

queritum 

Biosteres bevisi A Braconid parasite from South Africa (Wharton & Gilstrap, 1983) 

Trirhithrum sp. Rhynchosteres 

mandibularis 

A Braconid pupal parasite from Kenya ( Kimani-Njogu & Wharton, 2002) 

Zonosemata Biosteres sanguineus A Braconid parasite from Maryland to Florida to Missouri and Arizona 

electa 

(Wharton & Marsh, 1978). 

Zonosemata Diachasmimorpha A Braconid parasite from Minnesota and Nova Scotia to Florida (Wharton & 

vittigera 

(=Biosteres) 

sanguinea 

Marsh, 1978; Ovruski, et al., 2000). 


Natural predation, aside from micro-organisms, consist of birds, small 

animals and various invertebrates. While such predation is unlikely 

to influence outbreak populations of a fruit fly, there is some evidence 

that birds, ants, small mammals and other generalist predators are 

very important in suppressing fruit fly populations when the latter are 

already scarce and do act as a check under normal conditions 

(ie, Thomas, 1993). 

Bird Predation 

Should it develop that a resident bird population will effectively reduce 

the numbers of a targeted pest, then the bird population in question 

should be disturbed as little as possible. If it is felt desirable, the birds 

can be encouraged to increase in numbers through provision of food 

during winter months, the protection of nesting sites, and the 

discouragement of various bird predators or possibly, control of 

diseases. 

Small Mammal Predation 

Small mammals frequently prey on late instars and pupae and can 

remove large proportions of these individuals from a population. 

Pupae at or near ground level tend to suffer greater losses. 

Small mammals which feed on fruit flies include the white-footed 

mouse (Peromyscus leucopus) and the brush mouse (Peromyscus 

boylii). (Thomas, 1993) 


PPQ



Small mammals which are known or observed to feed on fruit fly life 

stages can be protected by not destroying their habitat or reducing 

their numbers through hunting. 

Insect Predation 

There is apparently an inverse relationship between vertebrate and 

invertebrate predation levels. Pupal predation by vertebrates 

increases as small mammal density increases, but invertebrate 

predation decreases (Cook, et al., 1995). 

See Thomas (1995), who stated that ants, rove beetles, and spiders 

were found to prey on Mexican fruit fly larvae which have left the host 

fruit and have not reached the shelter of a pupariation site. The 

predation rate was estimated to be 1-5%. Honey ants were recorded 

as being incidental predators. Rove beetles were deliberate predators 

during summer months. 

However, pupae and emerging adults were subject to attack by salticid 

spiders, an insectivorous ant, Pheidole sp. and especially exotic fire 

ants, Solenopsis invicta, which were effective predators. 

Approximately 29.5% of the fruit fly population was destroyed (70.5% 

survival). This factor was most effective in the summer. 

Ants may attack fruit fly last instar larvae on the ground as the latter 

look for a place to pupate. Ant numbers may be increased by spraying 

hosts with surcose; by encouraging benign (to hosts), host dwelling 

honey-dew producing aphids; by providing food for ants during fruit 

fly off season periods, or even by transporting ant nests into an area 

on a small scale (Weseloh, 1994). 

Similar results are known for other species of fruit fly. Ants 

(unspecified spp.) are responsible for 10% of the mortality of the 

soil-inhabitating stages of Bactrocera tryoni in Australia (Bateman, 

1968). In Hawaii, Pheidole megacephala attacks Bactrocera dorsalis 

larvae in fallen fruit, causing 36% mortality (Newell and Haramoto, 

1968). 

The interaction of ant species and prey may be summarized by stating 

that ant species occurring in an area are important agents in the 

regulation of pests and the rate of control varies spatially (ground, 

bush, tree) and temporally (winter, spring, summer, fall) within that 

area, depending on which species of ant is dominant. The dominance 

pattern is typically a mosaic of territories, as ant colonies partition the 

area in accordance with habitat requirements and competitive abilities 

(generalized from Thomas, 1995). Ant colonies may be encouraged in 

commercial areas by as little use of insecticides as is possible and 

covering the soil in a grass sward (Bateman, 1968). 


PPQ


Augmentation of Parasites/Predators - TABLE 3 

Spiders are another group of generalist predators that often consume 

the most abundant and most easily captured prey in their habitat. 

Encouragement of spider populations at present consist of not 

disturbing them; or of observing which species may feed on the larvae 

and bringing in more of these spiders from elsewhere to feed on the 

target fruit flies. 

Patch Complex 

A variation of the above, involves the employment of patch complexes, 

in which a number of areas are set up inside the entire control area to 

promote certain ecological situations advantageous for control within 

the economic constraints of a program. Inside the patch (or area), a 

complex of increased natural diversity is encouraged. Methods 

include the introduction of understory trees or bushes to increase the 

provision of nesting sites for birds, the encouragement/introduction of 

ant colonies, and so forth. 


Various efforts have been made to suppress fruit fly populations 

through augmentation of their parasites/predators. This technique is 

useful where more traditional methods such as insecticide-bait sprays 

and sterile male releases are impractical. The following table 

illustrates past procedures: 


PPQ




PPQ



Parasite/Predator Augmentation by Species—Target Pest - TABLE 3 

Species 

PARASITE/ 

PREDATOR RELEASE RATES RESULTS 

Anastrepha sp. Coptura haywardi ? Attacks pupae up to 5 cm deep. 

Suggested as a viable candidate to 

replace the environmentally 

unfriendly P. vindemiae (Guillen, et 

al., 2002) 


vindemiae 

Anastrepha ludens Diachasmimorpha 

longicaudata 

Anastrepha obliqua ? 

? 


longicaudata 

Anastrepha suspensa Diachasmimorpha 

longicaudata 

Anastrepha suspensa 

(Cont.) 


longicaudata 

? Attacks pupae on surface only 

(Guillen, et al., 2002). 

200,000 adult 

parasites/ week over 

100 ha in Mexico 

(Gingrich, 1993, citing 

unpublished data) 

940 adult parasites/ 

week in test plot in 

Mexico. 

? 

? 

940 adult parasites/ 

week in test plot in 

Mexico. 

60,000 adult 

parasitoids/km2 week 

by gallon buckets in 

hosts. NOTE: Fly larvae 

irradiated prior to 

parasitoid exposure to 

prevent a mixture of 

fertile flies/ parasites 

being release 

19,000-20,000 

parasitoids were 

released biweekly in 

defined 2.6 or 5.2 km 2 

release areas over 104 

km 2 . Release sites 

relocated to coincide 

with host availability. 

Release sites added 

when enough parasites 

were available to 

provide continuous 

linear release 

Apparently successful in Mexico & 

Costa Rica (Aluja, 1994, citing 

unpublished data) 

Suppression achieved 70% 

population reduction (Montoya, et 

al., 2000). 

Apparently successful in Mexico & 

Costa Rica 

(Aluja, 1994, citing unpublished 

data) 

Suppression achieved 70% 

population reduction (Montoya, et 

al., 2000). 

Target populations down to 5-10%. 

A second introduced parasite, 

Doryctobracon areolatus, nearly 

eliminated from release sit 


Target populations down to 4-21% 

(-95 & -79%) after augmented 

releases. (Burns, et al., 1996) 


PPQ

Parasite/Predator Augmentation by Species—Target Pest (continued)- TABLE 3 

Species 

Bactrocera dorsalis Diachasmimorpha 

longicaudata 

PARASITE/ 



tryoni 

1,694 o+ 212 o-> in 6 

hectare guava grove in 

1988 at the rate of 1-5 

times each month and 

25-90 o+ , 5-10 o-> 

each realease 

22,578 puparia in a 

100x140 meter open 

field or guava orchard 

by realease platforms 




by release platforms 

Bactrocera oleae Opius concolor Approx. 300-400 adult 

parasites per tree. 

Lower number = higher 

parasitism in upper 

reaches of tree. Higher 

number = overall 

parasitism throughout. 

Limited to target trees 

in middle of mature 

olive groves. Totals of 

2,500 and 4,000 O. 

concolor and 5,000 O. 

var. siculus released. 

Opius concolor var. 

siculus 

In Spain, 600 parasites 

per tree from August to 

October. 



Target populations down by 6 to 7% 

(Yao, 1989) 

Not Stated (Messing, et al.,1993) 

Not Stated (Messing, et al.,1993) 

Parasitism rate of 3rd stage larvae 

over 30%. Parasites can locate fly 

larvae efficiently on tre (Kapatos, et 

al., 1977) 

Flies decreased from 6.3 to 0.9 per 

trap (Gingrich, 1993, citing pers. 

com.) 


PPQ


Parasite Releases 

Parasite/Predator Augmentation by Species—Target Pest (continued)- TABLE 3 

Species 

Ceratitis capitata Diachasmimorpha 

tryoni 

Ceratitis capitata, Bactocera 

dorsalis, Bactrocera 

cucurbitae, Bactrocerus 

latifrons 

PARASITE/ 



longicaudata 


tryoni 

272,000 parasitized 

pupa/ week in 13 km 2 

area by gallon buckets 

in hosts in Hawaii. 

387,748 parasitized 

pupa/week in 13 km 2 

area by gallon bucket in 

hosts plus 3 million 

sterile Medfly by both 

aerial/ground releases 

in Hawaii. 




by realease platforms in 

Hawaii. 

Aerial releases of 

chilled adults in 

mountainous territory at 

286 to 846 parasites 

per hectare per week in 

Guatemala. 




by realease platforms in 

Hawaii 

Releases of various 

specific parasites for 

each species at a time 

for eradication from 

Hawaii. 

Rhagoletis cerasi Coptera occidentalis Thousands released per 

week under cherry trees 

in Czechosloviakia. 

Target populations significantly 

lower, parasitism to 47% from 

14.2%. No effect on parasitism of 

Eutreta xanthochaet (Wong, et al., 

1991). 

Target populations much lower, 

down to 9.8 Medfly adults/kg of 

fruit than in parasitic releases alone 

at 92.6 adults/kg of fruit (Wong, et 

al., 1992) 

Not Stated (Messing, et al., 1993) 

At higher release rates, parasitism 

levels reached 84% (Sivinski, et al., 

2000a). 

Not Stated (Messing, et al., 1993) 

Pilot trial against Ceratitis capitata 

in a 13 km sq area in 1982. 

(Gingrich, 1993) 

Parasitism ranged from 20-35% 

(Gingrich, 1993, citing personal 

com.) 


Releases are either made aerially or by static release stations on the 

ground. 


PPQ



Sivinski, et al. (2000a) demonstrated an aerial release method which 

relies on chilled adults in paper bags, these released from attitudes of 

100 meters at airspeeds of 130 km/hr. This is highly efficient and 

suitable for almost any type of terrain. 

Messing (1993) demonstrated a release platform that seems superior 

to the standard white plastic bucket release bucket of 3.5 or 4.5 liter 

capacity with escape holes just under the rim of the lid. The release 

platform is a circular release platform constructed to hold and protect 

the paper bags with parasitoids and allow the parasitoids to escape at 

any angle through a hardware cloth (see figure below). Use of the 

platform substantially reduced (by 29%) the mortality of parasitoids 

that died in the buckets. 

Augmentative, even undulative releases may not always work because 

of little known or poorly understood ecosystem constraints. If the 

parasites are released in ecosystems inhabited by an abundant and 

competitively superior parasitoid, the result may be little or no impact 

on host numbers. The size of the fruit may prove to be a limiting 

factor, as may encapsulation of parasite eggs by the host fruit fly. 

Then, too there is the possibility that the more efficient parasite may 

have already attacked the host, making it more sluggish and thus 

more prone to subsequent attack by the released parasite. Such a 

competitive situation could result in attack by the older parasite. 

(Purcell, et al., 1998) 


PPQ


Acknowledgements 

Acknowledgements 

FIGURE 3-4-1: hardware cloth = 1.2 cm 2 openings, height 10 cm, circumference 105 

cm Total escape area 1050 cm 2 

Grateful acknowledgements must be made to Dr. Dale Meyerdirk, 

USDA, APHIS, who first suggested that this information be made 

available to a wider audience than considered up to this point. The 

assistance of Ms. Rebecca Frasure of USDA, APHIS, PPQ, Manuals 

Unit, in the preparation of the manuscript, and particularly for the 

design and simplification of the Tables is also gratefully acknowledged. 

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