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Diptera: Ephydridae) for the Biological Control of the Submersed Aquatic Plant Hydrilla Verticillata (Hydrocharitaceae) in the Southeastern United States

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Diptera: Ephydridae) for the Biological Control of the Submersed Aquatic Plant Hydrilla Verticillata (Hydrocharitaceae) in the Southeastern United States BIOLOGICAL CONTROL 8, 65–73 (1997) ARTICLE NO. BC960491 Establishment of Hydrellia pakistanae (Diptera: Ephydridae) for the Biological Control of the Submersed Aquatic Plant Hydrilla verticillata (Hydrocharitaceae) in the Southeastern United States TED D. CENTER,* MICHAEL J. GRODOWITZ,† ALFRED F. C OFRANCESCO,† GREG JUBINSKY,‡ EDWARD SNODDY,§ AND JAN E. FREEDMAN† *USDA-ARS, Aquatic Plant Control Research Unit, 3205 College Avenue, Fort Lauderdale, Florida 33314; †USAE Waterways Experiment Station, 3909 Halls Ferry Road, Vicksburg, Mississippi; ‡Florida Department of Environmental Protection, Bureau of Aquatic Plant Control, 3917 Commonwealth Boulevard, Tallahassee, Florida 32399; §Tennessee Valley Authority, OSA 1B, Muscle Shoals, Alabama 35611 Received May 24, 1996; accepted October 8, 1996 Control of hydrilla is currently achieved primarily The Indian leaf-mining fly Hydrellia pakistanae De- with herbicides or by the release of exotic herbivorous onier was released in the United States during October fish (Sutton and Vandiver, 1986), but both methods are 1987 as a potential agent for the biological control of expensive for widespread use. Development of alterna- the submersed aquatic plant, Hydrilla verticillata (L.f.) tive control methods has been assigned high priority by Royle. Since then, nearly 3 million individuals have various federal and state agencies. One possible man- been released in four states at over 25 separate loca- agement alternative involves ‘‘inoculative’’ or ‘‘classi- tions. Hydrellia pakistanae is now permanently estab- cal’’ biological control, i.e., the introduction of host- lished at many locations in Florida as well as other specific plant-feeding insects obtained from within the portions of the southeastern United States. Despite native range of hydrilla. Searches for potential biologi- early failures, populations established from at least cal control agents began during the 1970s in India, 70% of the release attempts. Modifications of rearing Pakistan, and Africa and resumed during the 1980s in and release procedures that considered the biology of other areas of the world (Baloch and Sana-Ullah, 1974; the agent in relation to various habitat characteristics Baloch et al., 1980; Sankaran and Rao, 1972; Pember- facilitated successful establishment. 1997 Academic Press r ton, 1980; Center et al., 1990). The resultant surveys KEY WORDS: aquatic weeds; hydrilla control; biologi- revealed the presence of several biological control candi- cal control of weeds; Hydrellia pakistanae; Hydrilla dates including an undescribed Hydrellia species later verticillata; phytophagous insects; bioagent release named H. pakistanae Deonier (Deonier, 1978). techniques; Ephydridae; Hydrocharitaceae. Hydrellia pakistanae is a small, ephydrid fly whose larvae mine hydrilla leaves. Eggs are laid on floating leaves and the larvae either enter the leaves directly or INTRODUCTION descend into the water column before entering the plant. Females apparently oviposit indiscriminately, Hydrilla, Hydrilla verticillata (L.f.) Royle (Hydro- but the larvae are quite selective (Buckingham et al., charitaceae), is a submersed, leafy-stemmed vascular 1989; Wheeler and Center, 1996). Each larva mines hydrophyte that is widely distributed in the Old World. about 12 leaves during the course of its development It was introduced into the United States through the and then pupates within an air-filled puparium that is aquarium trade in the early 1950s (Schmitz et al., attached in a leaf axil (Baloch and Sana-Ullah, 1974; 1991). Currently, infestations of this invasive plant Buckingham et al., 1989). Total generation time is 18 to constitute the most severe aquatic plant problem in the 30 days (Baloch and Sana-Ullah, 1974). southern United States. Hydrilla infested 22,000 ha in Laboratory studies done in Pakistan (Baloch et al., Florida by 1988 and the state was spending about $7 1980) and later in a U.S. quarantine facility located in million annually in attempts to control about 6000 ha Gainesville, Florida (Buckingham et al., 1989), verified (Schmitz et al., 1991). During this same period it that H. pakistanae was host-specific to hydrilla. Accord- rapidly expanded its range and now occurs as far west ingly, Dr. Gary Buckingham petitioned the USDA Ani- as California (Sonder, 1979). mal and Plant Health Inspection Service—Plant Protec- 65 1049-9644/97 $25.00 Copyright r 1997 by Academic Press All rights of reproduction in any form reserved. 66 CENTER ET AL. tion and Quarantine (USDA-APHIS-PPQ) in June 1987 ther separately or in mixture (4 g yeast hydrolysate for permission to release this species. Permission was plus 7 g sucrose in 7 ml of water). Partially submersed obtained after review by the Technical Advisory Group hydrilla sprigs placed in shallow dishes within the on Biological Control of Weeds (Coulson, 1992) and this chambers provided the oviposition substrate. Egg- insect was first released on 29 October 1987 (Bucking- laden sprigs were transferred to new water-filled con- ham, 1988). In this paper, we document the results of tainers after 1 to 3 days to perpetuate the rearing this and later releases of H. pakistanae and the subse- process. quent establishment of self-perpetuating field popula- The second rearing method consisted of placing tions. This project represents the first attempt to use an freshly harvested hydrilla in large tanks. Either adults insect to control a weed in the submersed aquatic or hydrilla sprigs containing immatures were placed on environment by implementation of the ‘‘classical’’ ap- the floating hydrilla. Fly populations were allowed to proach to biological control. increase in size within the tanks without additional handling. The hydrilla was then removed and trans- ferred to field sites. MATERIALS AND METHODS Releases. The basic release method involved the Insects. Buckingham et al. (1989) documented the placement of insects directly on hydrilla infestations collection of the original H. pakistanae stock from India either within open hydrilla beds or in protective cages and its receipt and disposition at the Gainesville, (Fig. 2). The cages were constructed of two frames made 2 Florida, quarantine facility during May to August 1985. of 2.5 cm (cross section), channeled aluminum extru- This initial stock was used for all host specificity sion rods. The frames were attached to one another testing and provided progeny for the initial U.S. re- with a piano hinge. The tops of the upper frames were leases made during 1987 to 1990. However, because of covered with fine-mesh screen (different materials with the risk associated with excessive inbreeding of the varying mesh sizes were used but 60% greenhouse original stock, fresh material was obtained from over- shade cloth usually sufficed). Floatation was provided seas. The Gainesville quarantine facility received a by filling the hollow aluminum extrusion rods with fresh consignment of field-collected H. pakistanae from polystyrene. The cages were placed at each site on the Kumbaleagodu, Bangalore, Karnataka State, India, on water surface with the screened side up. To prevent 25 August 1990 and one from Rawalpindi, Pakistan, on lateral movement, the cage was anchored by passing 28 August 1990. These insects were reared in quaran- poles through U-bolts attached to the outside of the tine for less than three generations prior to field release lower frame. The ends of the poles were forced into the to confirm identifications and to eliminate parasites. bottom soil. The hinged upper frame provided access After permission was granted for release of H. paki- into the cage for fly releases and for later assessments. stanae, a portion of the original quarantine colony was Numerous variations of the basic release technique used for the first release, a portion was retained in were employed. For example, individuals from both quarantine, and a portion was transferred to the U.S. laboratory colonies and from previously established Department of Agriculture, Agricultural Research Ser- field sites were released. Most of the individuals from vice (ARS), Aquatic Plant Control Research Unit in laboratory colonies were progeny of the original quaran- Fort Lauderdale, Florida. Insects reared at ARS were tine colony. However, some freshly acquired material later provided to the Waterways Experiment Station from overseas was released at selected sites. All life (WES) of the U.S. Army Corps of Engineers in Vicks- stages were released although the majority were eggs burg, Mississippi, the Tennessee ValleyAuthority (TVA) or larvae. Immature stages were transferred on or in Muscles Shoals, Alabama, and to the Florida Depart- within the plant tissue to minimize mortality. ment of Natural Resources (FDNR, now the Depart- The number of releases made and the total number of ment of Environmental Protection) Bureau of Aquatic individuals released varied among sites. In some cases, Plant Control in Tallahassee, Florida. predetermined quantities were released. In other in- Rearing. Two basic techniques were used to rear stances, releases were made repeatedly and were discon- H. pakistanae. The primary method was modified tinued only after evidence of establishment was ob- slightly from that described by Buckingham et al. tained. The brief growing season limited the numbers (1989). Hydrilla sprigs containing freshly oviposited released at more northern sites (i.e., Alabama). eggs were placed in 2- to 4-liter water-filled containers Assessment of establishment. Sites were checked for containing approximately
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