Progress 01/01/04 to 12/31/04
Outputs
The work completed in 2004 was done in the context of the four primary objectives of this project: 1) diagnostic DNA tools for maggot identification; 2) biological control of tephritid pests; 3) traditional taxonomic work on tephritid flies; and 4) host associations of flies and parasitoids in field-infested fruits. An important side-benefit of the project is that seeds from rare plants, or those species that have been under heavy predation pressure due to logging, were saved following the rearing of insects from fruits and given to Kenyan botanists and foresters for propagation. Objective 1 has been largely completed. PhD student Norman Barr completed his dissertation under collaborator McPheron at Penn State. The research included development of a diagnostic tool for identification of ceratitidine tephritids, and we had an opportunity to test this on maggots that were intercepted from a shipment originating in Europe. ICIPE collaborators Osir and Masiga also
finished their portion of the work, with Osir co-authoring one paper. Additional collections were made in Kenya to help resolve problem taxa in the rosa-fasciventris group. All laboratory studies associated with objective 2 were completed by the end of the year. This included work on 3 species of Psyttalia and on Tetrastichus giffardii by S. Mohamed. Identifications were provided for biocontrol programs on olive fly, and we made several attempts to collect more olive fly parasitoids in Kenya. Our website on parasitoids of Tephritidae was updated. Two African PhD students completed their theses under this objective. Objective 3 included the incorporation of several thousand additional specimens into the web-accessible, Afrotropical tephritid database (M. De Meyer and I. White) and the revision of Afrotropical dacines (White). British Museum collaborator White has a conventional key now available, as well as a preliminary version of a CABIKEY product and has compiled 1300 images for
this work. He is now looking for a publication outlet for the revision which is several hundred pages long. As in previous years, workshops were conducted on identification of pest species, mostly in Europe. Additional work was done on the new invasive species noted last year, and a tentative identification has now been made of this mango pest. Under objective 4, a paper on flies and parasitoids developing in fruits of Oleaceae was published and several additional collections of wild fruits were made while searching for olive fly parasitoids. This project continues to provide numerous opportunities for collaboration with scientists from around the world. In addition to olive fly work noted above (collaborators from France, South Africa, Italy, Hawaii, and California), we are co-authoring publications on on pest tephritids and their natural enemies with Clarke and Carmichael in Australia, Aluja, Rull, and Ovruski in Mexico and Argentina, and Ngoume from Nigeria, and recently signed a
cooperative agreement with Vargas at the USDA/ARS lab in Hawaii.
Impacts
Major impacts were in the area of identification of fruit-infesting tephritid pests and their natural enemies. Collaborators White and De Meyer provided identifications of the newly introduced exotic mango pest, enabling workers to trace the spread of this species throughout eastern Africa and eventually to western Africa. Thanks to the work of White during the course of this project, we are now able to provide identifications for virtually all Afrotropical dacines, which has been a long-standing problem. Another long-standing problem has been the identification of immature stages in infested fruits. This has now been solved for a large group of pests and potential pests through the development of DNA-based diagnostic tools by collaborators Barr and McPheron. Of special interest was the development of a non-destructive extraction protocol for DNA, which will considerably facilitate all future work in this area. Barr also discovered that one of the more commonly
encountered species in tropical Africa consists of several discrete populations that can be recognized by distinct DNA signatures. As in other years, Wharton provided routine identifications of numerous parasitoids, and supplied extensive information on the parasitoids of olive fly relative to biological control efforts against this species. Much of this information has now been transferred to web-accessible format to facilitate use by applied entomologists. Our website on parasitic Hymenoptera at Texas A&M was chosen for inclusion in Natural Selection (co-ordinated in part by the The Natural History Museum, London).
Publications
- Mwatawala, M.W., White, I.M., Maerere, A.P., Sendondo, F.J., and De Meyer, M. 2004. A new invasive Bactrocera species (Diptera: Tephritidae) in Tanzania. African Entomol. 12:154-156.
- Copeland, R., White, I.M., Okumu, M., Machera, P. and Wharton, R.A.. 2004. Insects associated with the fruits of the Oleaceae (Asteridae, Lamiales) in Kenya, with special reference to the Tephritidae (Diptera). Bishop Museum Bull. Entomol. 12:135-164.
- Barr, N.B. 2004. The molecular systematics of Ceratitis Macleay (Diptera: Tephritidae). Ph.D. dissertation, Pennsylvania State University.
- Baliraine, F.N., Bonizzoni, M., Guglielmino, C.R., Osir, E., Lux, S.A., Mulaa, F.J., Gomulski, L.M., Zheng, L., Quilici, S., Gasperi, G. and Malacrida, A.R. 2004. Population genetics of the potentially invasive African fruit fly species, Ceratitis rosa and Ceratitis fasciventris (Diptera: Tephritidae). Mol. Ecol. 13: 683-695.
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Progress 09/15/00 to 09/30/04
Outputs
All four primary objectives (development of diagnostic DNA tools for maggot identification; selection of natural enemies for biological control of tephritid pests; taxonomic work on tephritid flies; and discerning patterns of utilization wild host plants by frugivorous tephritid pests) were accomplished. Thus far, we have published or submitted 22 peer-reviewed articles based on the work done under the primary objectives. Four dissertations were also completed on research supported entirely or in part by this grant. Publications not included in previous reports are listed below. Additionally, several studies based on non-target insects reared from our fruit samples have been published or are in preparation, one cd has been developed on Kenyan fruits and their associated fauna, and at least 8 presentations on our work have been made at national and international meetings. Specific outreach components include the work of Heather Ketchum at Texas A&M University, who
developed a hands-on curriculum for investigating the impact of invasive species on native flora and fauna, with different modules that cover grades 4-12. The PI also conducted a graduate student seminar on invasive species during the first year of the project. Many of our publications are taxonomic, addressing both species-level and genus-level problems that hinder pest identification. Among the more significant of these is the segregation of members of the Ceratitis rosa species group and clarification of the identities of cucurbit-infesting species in the genus Dacus. Several workshops on pest and parasitoid identifications were conducted by White, De Meyer, and the PI, largely directed to pest management personnel. Hands-on training was provided to scientists and technicians from at least 14 countries, in addition to large numbers of identifications (and associated information) for both flies and parasitoids provided to fruit fly workers in over 30 countries, mostly from the third
world. To facilitate transfer of information to applied entomologists in the future, on-line databases and web-based keys have been developed. The multiple entry key to Dacina, for example, will have approximately 1300 images. Preliminary databases developed under this grant were used to acquire funding from European sources for expanding the work of De Meyer, White, and Mansell on specimen-level databases for tephritid fruit flies. Major collections of pests and their close relatives, totalling several thousand completely curated and determined specimens, were deposited in major museums in five countries. Smaller, synoptic collections for use in detection programs were deposited in Florida and California, the two most important states for interception of tropical and subtropical fruit pests. Biological control programs on medfly, Natal, fly, olive fly, and cucurbit flies benefited by the collection of several parasitoid species, including work by graduate students on culture methods,
host-range testing, and ultimately the shipping of seven species to collaborators in Hawaii, California, Guatemala, France, and St. Helena.
Impacts
Our dissemination of information about fruit-infesting tephritid pests and their natural enemies has facilitated the efforts of state and federal agencies involved in detection, eradication, and management (including biological control) of invasive, frugivorous pests such as medfly, olive fly, Natal fly, and various cucurbit flies. Both traditional and electronic, image-intensive publications have simplified pest and natural enemy identifications, and the first significant diagnostic tool for Ceratitis larvae has been developed and successfully tested. The latter enables rapid identification of larvae in infested fruits during monitoring and detection programs, so that appropriate, species-specific protocols can be activated. Lists of susceptible hosts, widely used in tephritid pest management, were significantly expanded for all of the pest species that we sampled. These data have been published and made available in web-accessible databases, together with detailed
annotations on previous host records. Our data provide some of the first assessments of patterns of host plant utilization by widespread tephritid pests in their area of endemicity, and support an East African origin of the medfly. The intensive collecting effort also provided baseline data for East Africa that enabled immediate detection of a newly invasive mango pest. New species of tephritids and their parasitoids were discovered and described, and potential natural enemies of tephritid pests were reared, cultured, evaluated, and shipped to collaborating scientists for use in biological control programs.
Publications
- Amy E Carmichael, A. E., Robert A Wharton, R. A., and Clarke, A. R. 2005. Opiine (Hymenoptera: Braconidae: Opiinae) parasitoids of tropical fruit flies (Diptera: Tephritidae: Dacinae) of the South Pacific region. Accepted, Bull. Ent. Res.
- Ovruski, S. M., Wharton, R. A., Schliserman, P., Aluja, M. 2005. Abundance of Anastrepha fraterculus (Diptera: Tephritidae) and its Associated Native Parasitoids (Hymenoptera) in Feral Guavas Growing in the Endangered Northernmost Yungas Forests of Argentina with an Update on the Taxonomic Status of Opiine Parasitoids Previously Reported in this Country. Accepted for publication, Ann. Entomol. Soc. Am.
- Copeland, R. S. and Wharton, R. A. 2005. Year-round production of pest Ceratitis species (Diptera: Tephritidae) in fruits of the invasive species Solanum mauritianum in Kenya. Submitted to Ann. Entomol. Soc. Am.
- Copeland, R. S., Wharton, R. A., Luke, Q., De Meyer, M., Lux, S., Zenz, N., Machera, P., and Okumu, M. 2005. Geographic distribution, host fruits, and parasitoids of the African fruit fly pests Ceratitis anonae, Ceratitis cosyra, Ceratitis fasciventris and Ceratitis rosa (Diptera: Tephritidae) in Kenya. Submitted to Ann. Entomol. Soc. Am.
- Barr, N. B., Copeland, R. S., De Meyer, M., Masiga, D., Gathenga, H., Osir, E., Robert Wharton, R. A., and Bruce A. McPheron, B. A. 2005. Molecular diagnostics of economically important fruit fly species (Diptera: Tephritidae) in Africa using PCR and RFLP analyses. Submitted to Bull. Ent. Res.
- Copeland, R. S., Okeka, W., Freidberg, A., Merz, B., White, I. M., De Meyer, M. and Luke, Q. 2005. Fruit flies (Diptera: Tephritidae) of Kakamega Forest, Kenya. Submitted to Journal E. Afr. Nat. Hist.
- De Meyer, M., Copeland, R. S., Wharton, R. A., and McPheron, B. A. 2004. On the geographic origin of the Medfly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Proc. 6th Intl Fruit Fly Symposium, Stellenbosch, pp. 45-53.
- Vayssieres, J.-F., Wharton, R., Delvare, G., Sanogo, F. 2004. Diversity and pest control of hymenopteran parasitoids of Ceratitis spp. on mangoes in Mali. Proc. 6th Intl Fruit Fly Symposium, Stellenbosch, pp. 461-464.
- Mohamed, S. 2004. Suitability and acceptability of five fruit infesting tephritids for three Psyttalia spp. parasitoids. Ph. D. Dissertation, University of Gezira. Wad Medani, Sudan.
- M. Billah. 2005. Biosystematic studies of Psyttalia species (Hymenoptera: Braconidae): Parasitoids attacking fruit-infesting flies (Diptera: Tephritidae) in Africa. University of Ghana
- Trostle, M. 2005. Classical biological control of Mediterranean fruit fly, Ceratitis capitata (Wiedemann), (Diptera: Tephritidae): natural enemy exploration and nontarget testing. Ph.D. Dissertation, Texas A&M University.
- DeMeyer, M. and Copeland, R. S. 2005. Description of new Ceratitis MacLeay (Diptera, Tephritidae) species from Africa. J. Nat. Hist. 39: 1283-1297.
- De Meyer, M. and Freidberg, A. 2005. Revision of the fruit fly genus Capparimyia (Diptera, Tephritidae). Zool. Scripta 34: 279-309.
- Freidberg, A. and Copeland, R. S. 2005. Notommima parallela, a new genus and species of fruit flies from Kenya allied to Notomma Bezzi (Diptera: Tephritidae). Submitted to J Nat. Hist.
- White, I. M. 2005. Taxonomy of the Dacina (Diptera: Tephritidae) of Africa and the Middle East. Submitted to Sth. Afr. Entomol.
- Lux, S. A., R. S. Copeland, I. M. White, A. Manrakhan, and M. K. Billah. 2003. A new invasive fruit fly species from the Bactrocera dorsalis (Hendel) Group Detected in East Africa. Insect Sci. Applic, 23: 355-361.
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Progress 01/01/03 to 12/31/03
Outputs
The project contains four basic components: 1) Diagnostic DNA tools for maggot identification; 2) biological control of tephritid pests; 3) traditional taxonomic work on tephritid flies; 4) host associations of flies and parasitoids in field-infested fruits. Additionally, as part of our outreach activities, a PhD student at Texas A&M (Heather Ketchum), using funding from this grant, developed lesson plans on invasive species for primary and secondary schools. For objective 1, graduate student Barr presented preliminary results at the Evolution Society meetings in June and Kenyan collaborator Masiga worked in McPheron's laboratory at Penn State to develop protocols for field testing results from McPheron's lab on known and unknown populations in Kenya. Work on objective 2 focused on completion of laboratory studies on host specificity and development of parasitoids of medfly, African mango fly, lesser pumpkin fly, and olive fly; cultures of olive fly parasitoids were
also established and shipments were sent to France under a cooperative agreement with USDA; the PI identified several lots of parasitoids for various state and federal agencies involved in biological control of olive fly in California; also under this objective, two PhD students were supervised, 3 papers were initiated, two others were submitted, and two more were published. Ian White completed a major portion of his Afrotropical Dacinae under objective 3, including a preliminary dichotomous key to African Dacini; Marc De Meyer continued to work on Ceratitis, including collaboration with White and Mervyn Mansell on web-accessible databases; both White and De Meyer continued to provide identifications and other information for Afrotropical tephritids as part of this program, and also conducted training programs for fruit fly identification; due to our on-going work in Kenya, White was able to determine that a newly discovered fly in mango was an invasive species not previously known in
Africa (USDA has now taken over the program for management of this newly invasive species); significant collections of determined species of Afrotropical Tephritidae (from our sampling program) were curated and delivered to the US National Museum, The British Museum of Natural History, and two museums in Kenya this year. Under objective 4, a paper on the tephritid flies of olives in Kenya, co-authored by White, postdoc Copeland, and others was submitted; three additional works on geographical distribution and native host plants of tephritid pests were either initiated or submitted; new fruit collections were made in northern Kenya, an area previously considered too dangerous to sample, and several new host plants were obtained. Thus far, the PI, co-investigators White and De Meyer, and postdoc Copeland have developed new collaborative ties to researchers from 9 additional countries as a direct result of this program (beyond the routine identifications of insects provided to a much
larger group). Numerous state and USDA scientists have also participated as co-authors of papers and recipients of training programs, shipments of natural enemies, and/or data essential to the conduct of their programs.
Impacts
The most significant impact of our program in 2003 was the discovery of a new invasive pest in East Africa. Had this project not existed, the newly invasive species might still be undetected. The species was first collected in an area where we have been sampling native host plants intensively to improve our understanding of patterns of host fruit utilization by fruit fly pests. Since our field work was done in close collaboration with taxonomic work, also supported under this project, we were the only research group in a position to provide the necessary information that this particular fly was indeed a new invasive pest. Additionally, the co-investigators from Belgium, England, Kenya, and 3 U.S. institutions worked with scientists and students from at least 14 other countries, providing training sessions, information and identifications of pest tephritids and their natural enemies used in biological control. In many cases, the information provided was a direct result
of this IFAFS-supported research, much of it still unpublished. Nearly all of this effort was above and beyond what each of us does individually in this regard; the IFAFS collaboration has thus considerably enhanced our contributions to pest detection and control internationally. The PI has also provided nearly all identifications (and associated biological information) for parasitoids associated with the biological control effort in California against olive fly, a new invasive species; and we are the only research group that has been able to culture one of the parasitoids, Psyttalia lounsburyi.
Publications
- M. Lopez, J. Sivinski, P. Rendon, T. Holler, K. Bloem, R. Copeland, M. Trostle, and M. Aluja. 2003. Colonization of Fopius ceratitivorus, a newly discovered African egg-pupal parasitoid (Hymenoptera: Braconidae) of Ceratitis capitata (Diptera: Tephritidae). Florida Entomologist 86: 53-60.
- Mohamed, S. A., W. A. Overholt, R. A. Wharton, S. A. Lux, Elameen, M. Eltoum. 2003. Host specificity of Psyttalia cosyrae (Hymenoptera: Braconidae) and the effect of different host species on parasitoid fitness. Biological Control 28: 155-163.
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Progress 01/01/02 to 12/31/02
Outputs
Efforts during the second year of this project were focused on three objectives: 1) sampling of pest tephritids and their natural enemies; 2) systematics research on flies and the parasitic wasps that attack them; and 3) biological control of pest tephritids of African origin. Sampling efforts for the entire project were largely completed this year. The sampling program was primarily designed to obtain fresh material of a diverse array of fruit-infesting tephritids for the second objective. The intense sampling effort in Kenya, coordinated by postdoctoral researcher Bob Copeland, also provided a wealth of data on regional and seasonal patterns of host plant use by these flies. Two papers on host plants of Afrotropical fruit-infesting tephritids were published this year and several more are now being written. The second objective has two parts. One is focused on traditional, descriptive taxonomy of the flies and their parasitoids, with the products ultimately
translated into electronic databases and user-friendly, multiple-entry, electronic keys. Dr. Ian White, a collaborator from England, has made considerable progress on the Afrotropical members of the genus Dacus, a large and difficult group. He and fellow collaborator Marc De Meyer, from Belgium, have also provided critical identification of the material that formed the basis of our published host records as well as all specimens sent to collaborator Bruce McPheron for the molecular diagnostic research. Molecular characterization of species was begun in earnest during this second year, and preliminary results show that some groups of pests are readily characterized by the relatively short strands of DNA that have been analyzed thus far. A few key pest species will clearly need more work, and different gene regions are now being explored. In addition to McPheron's efforts at Penn State, Ellie Osir and Dan Masiga have recently initiated their investigations of the population genetics of
3 species in Kenya, also using material sampled this year. We have also been facilitating the work of other systematists by sending them various other insects (e.g. moths, weevils, seed wasps) that we've reared from fruits. The third objective (biological control) was a major focus of this year's effort. Activities can be listed under three somewhat overlapping categories. First, most of the laboratory studies on various tephritid parasitoids were completed and several of these have been prepared for publication. These studies have focused primarily on basic life history traits of species in the genus Psyttalia and to a lesser extent Fopius and Diachasmimorpha. Graduate students Samira Mohamed and Marcia Trostle have conducted most of the research. Second, several parasitoid species were sent to colleagues in Guatemala and Hawaii where additional non-target studies will be conducted prior to possible field release for biological control of various pests. Third, direct shipments were
made to California and the island of St. Helena for immediate release (following quarantine clearance) against invasive species (olive fly, Bactrocera oleae and lesser pumpkin fly, Dacus ciliatus respectively).
Impacts
Direct impacts of our research include facilitation of biological control programs in the U.S., Guatemala, and St. Helena, primarily against invasive pests such as olive fly and medfly. We have characterized the biology of several natural enemies and shipped them to potential release sites. Publication of host records firmly establishes the polyphagous nature of medfly in its native home but also highlights the fact that it does have preferred hosts. Many old literature records of Afrotropical fruit-infesting flies have been corrected and updated.
Publications
- Copeland, R. S., R. A. Wharton, Q. Luke and M. De Meyer. 2002. Indigenous hosts of Ceratitis capitata (Diptera: Tephritidae) in Kenya. Annals of the Entomological Society of America 95: 672-694.
- De Meyer, M., R. S. Copeland, S. A. Lux, M. Mansell, S. Quilici, R. Wharton, I. M. White, and N. J. Zenz. 2002. Annotated check list of host plants for Afrotropical fruit flies (Diptera: Tephritidae) of the genus Ceratitis. Koninklijk Museum voor Midden-Afrika Tervuren, Belgie, Zoologische Documentatie 27: 1-91.
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Progress 01/01/01 to 12/31/01
Outputs
This project is multi-faceted. It includes a sampling component for tephritid fruit flies that includes both known and potential pests. There is also a systematics component that treats both flies and their natural enemies. The primary goal of the systematics component is to facilitate identification of pests when intercepted in commodities and to provide a sound classification of the parasitoid wasps that attack these pests. There is also an extension/outreach component to this project. The sampling component, conducted largely by Bob Copeland, has been on-going for a year, with excellent results. We have completed our survey of native hosts of Mediterranean fruit fly, Ceratitis capitata, one of our key targets from the standpoint of invasive potential affecting US agriculture. This work was conducted in Kenya (Medfly is native to east Africa), where fruits were sampled from diverse areas including lowland, tropical forests along the coasts, remnants of the
equatorial rainforests in western Kenya, semiarid scrub, and mountain habitats. Medfly was reared from 55 species of plants, 51 of them indigenous. Forty-five of these species represent previously unknown hosts in Africa. Important host plants were noted, as was the relatively uneven attack rate: some hosts are obviously preferred while others are never attacked. The results of this survey will shortly be submitted for publication. Similar surveys have been conducted for other known pest species, and these will be summarized for publication in the near future. In addition to providing excellent records on patterns of host fruit utilization, these collections have yielded an wealth of material that is now being used in the systematics research. The latter includes completion of a revision of the genus Ceratitis by collaborator Marc De Meyer of the Tervuren Museum, Belgium, initiation of a revision of the Afrotropical species of the genus Dacus by Ian White (British Musuem), studies on
opiine braconid parasitoids by Wharton, and (as one of the central features of this program), molecular characterization of species in the genus Ceratitis through a combined effort by Bruce McPheron at Penn State and Ellie Osir at ICIPE in Kenya. Several shipments of appropriately preserved material are now at Penn State as a result of the sampling program. We have also obtained material for the molecular studies from South Africa and Ghana. We have also initiated an outreach component that consists of a white paper on invasive species that was submitted to the Entomological Society of America and is currently under review. There is a biological control component that currently involves host range testing of parasitoids in Kenya and Hawaoo as well as an effort to release natural enemies against medfly, olive fly, and one of the indigenous curcurbit flies that is a widespread pest in Africa.
Impacts
The work by Ian White will enable accurate and confident identification of all Afrotropical Dacus pests for the first time. Currently, only a few of the species are sufficiently well-known. The main impact, however, will be the shortening of identification time for fruit infested with larvae, a key concern of quarantine personnel.
Publications
- No publications reported this period
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