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Suppl. No. 25 Southwestern SEPT.2OO2 SUPPL.NO. 25 SOUTHWESTERNENTOMOLOGIST PHORIDFLIES FORTHE BIOLOGICAL SUPPRESSIONOF IMPORTEDFIRE ANT IN TEXAS: REGIONSPECIFIC CHALLENGES' RECENT ADVANCES AND FUTUREPROSPECTS LawrenceE. Gilbertand Richard J.W. Patrock Sectionoflntegrative Biology andBrackenridge Field Laboratory TheUniversity of Texag,Austin 78712 ABSTRACT At the Universityof Texas'Brackenridge Field Laboratorywe maintainone of two laboratoriesin the U.S.devoted to researchon the basicbiology of Pseudacteonphorid flies' a diversegenus offire ant parasitoids.Goals include 1) screeningvarious South American specieso{Pseudacteon for their potential to negativelyimpact the red imported fue ant, S. i)victa,2) testing prospectspecies for their host specificity,3) developing.methodsfor rearingcandidatJspecies, and successfullyintroducing them to Texas,and 4) monitoring impact of fire ant phoridson local ant communityorganization. Challengeiin introducingtopical P. tricuspisto confol S. invicta populationsacross central and ,outh T.tut include the l) high rate ofpolygyny in S. irwicta populationsand 2) harsh,unpredictable climate of the region.Nevertheless, field-bred P. tricuspishave been ,""olr"r"d in 9 of 15 experimentalrelease sites over periods:upto27 months.In South America, we have collaboratedwith colleaguesto study the life histories, geography,and seasonalphenologies of little known Pseudacteonspecies such as P. cultellatus.Our laboratory experirnentshave shownthat different speciesof Pseudacteonvary in the extent to which they disrupt the foraging activities of fire ants. Laboratory experimentsrevealed the negativeimpaciof P. tricuspison colony protein consumptionwhich ultimately affects colony-fitness.bur resultsfrom variousinvestigations lead to the conclusionthat in Texas, biologicalcontrol of fire antsusing phorid flies will likely requiremore than a singlespecies of Pseudacteon,as well asregion-specific tactics. INTRODUCTION The associationof phorid flies of the genusPseudacteon with fire antshas been known sincew.M. wheelei and c.T. Bruesstudied the biology of solenopsisgeminata in central Texas early in the 20thcentury (Coquillett 1907;Brues 1907).Until the 1990s, researchon pseudacteonwas primarily taxonomic, but their associationwith disturbednests ofthe genusSolenopsis was notedin the descriptionsofmany species.Efforts in the early 1970'ito find biologicalcontrols for SolenopsisinvictaBven (Hymenoptera:Formicidae) in South America identifiedPseudacteon among other possibleagents (Williams 1973; Williams 1974).While thesephorids were subsequently considered possible candidates for biologicalconirol of S. invicta (Williams et.al.7973), efforts were not sufficientto assess the dJtailsconceming phorid/ant interactions. From the perspectivethat soughtbiological agentsthat imposedhigi mortalityrates on targetpest species, Pseudacteon species showed lilttle appa.enipromis- and were not pursuedby thoseresearchers' The work of Feener (1981)stimulated interest in Pseudacteonphorids as potential biological control agents for fire ants. Studyingcompetition between Pheidole dentataMayr and Solenopsistexana Emeryat BrackenridgeField Laboratory(BFL) in Austin Texas,Feener demonstrated that the presenceof phoridswhich attackP. dentatamajors shifted the outcomeof competitive interactionswithStexana. Withoutphorids, Pheidole always won controlof food baits.With phorids Solenopsiswon. Feener'sdemonstration of the behavioraleffects that phorids imposeon ant workers indicatedthat the primary impactsof theseparasitoids on ant colony fitnessshould be indirectvia reducingfood intake(especially in the contextofcompetition with other organisms)rather than by the direct mortality of workers. Feener'shypothesis, extended to importedfire ants,predicted that an important differencebetween native non-pest S. geminata F. andthe invasivepest S. invictais thatthe latterhas escaped its Pseudacteonfawa. Morrison(2000a;2000b) has recently shown the way that Pseudacteonspecific to s. geminata(P. browni Disney,P. biJidusBrown and Morrisorl P. spatulatusMalloch) reducethat species'ability to competefor food with S. invicta in Texas.Does freedom from phorid pressurein its introducedrange account for the ability of s. invictato replaces. geminata(e.g., Porter et al. 1988)?In Brazil,pseudacteon often alter the ability of S. invicta to successfullycompete for food resourcesduring the day (On et al. 1995;Porter et al. 1995). Theseand other observationsand experimentsunderlie the working assumptionsof our researchprogrtlm: l) lhat Pseudacteonphoids ultimatelyaffect the populationlevels and ecologicaldominance ofhost fire ants,and 2) that theseparasitoids are a major factor influencing the population and community biology of the North American Geminata and SouthAmeric an Saevissimaspecies complexes of the genusSolenopsis, The latter species complexincludes S. invicta and morethan a dozenrelated species native to SouthAmerica. In fall 2000,usDA APHIS announcedan expandedeffort to fund the productioilof Pseudacteontricuspis Borgmeier in partnershipwith ARS (ARS News Service,Novenlber 15,2000) for releasetlroughout the southemuS. This newsrepresented a tuming poinf in terms of official recognitionof the potential of biological control of fire ants. More specifically,it identifiedPseudacteon phorids as oneofthe leadingprospects for reducing the peststatus of,Solezopsis richteri Foreland S. invicta.Furthermore the burden of rearing P. tricuspis,by that decision,was transferredfrom the only phorid researchlaboratory within the USDA, thus allowing it to allocatemore attentionto fundamentalresearch on less well-knownPseudacteon species and their relationships with fire ants. The USDA's commitmentto mass-rearand releasep. tricuspis,the first phorid species(of many possiblealtematives) brought into cultivation,should be viewed as a positiveinitial step.It is difficult to know at this earlystage however, ifthis is the besfstep possible.It is alsounclear whether Brazilian P. tricuspiswill be effectivein Texas.Bqlow, we discusspotential problems offocusing on a singlespecies ofphorid fly for biological control ofthe red imported fire ant over the large geographicalrange in the United Siates now occupiedby this ant. REGIONSPECIFIC CHALLENGES FoR THE usE oF pseudacteontricuspis There are compellingarguments for separate,but cooperatingefforts to develop different phorids for fire ant suppression.First, the biogeographicpatterns and eco- genotypic diversity within the genls Pseudacteonare relativeiy unexploredsubjects. second,the rangenow occupiedby invadings. invicta in the US is highiy diverse,both edaphicallyand climatically.In fact, additionalresearch teams are neeaiaio complement thosein Florida and Texasif we are to rapidly explorethe full potentialor phoiid flies acrossa diverseecological template, which now includesparts ofArizona andCilifornia. - Two region-specific-challenges,important when consideringthe applicationof phoridflies to suppressionof fire antsin Texas,are special foci of ouiresearch-group. The first major challengeis the harshnessand unpredictabilityof the Texas climate. Both Gainesville,FL and Austin,TX laboratorycultures of P. tricuspisare based on collections originatingfrom severallocal sitesfrom SdoPaulo State Brazil, a tropicalregion with dry and wet seasons.Our initial field releaseof P. tricuspisat BFL in November1995 was followedby a harshdrought which lastedinto August1996. In 1996,February temperatures beganat -9' C andended at 37" C; extrememonthly records for all Austinweather stafions between1930-2001 (NOAA 2001).Exceedingly arid andwarm springand fall periodshave beena frequentfeature of Texasclimate during the courseof our work on phoridreleases in Texas(1996-2000). Palmer drought indices place these years as being one ofthe threedriest periodssince records have been kept (NOAA 2001).Such dry conditionsduring the periods when temperaturesare generally favorable for tropical phorids make successful establishmentof releasedpopulations more difficult. Ironically, during theserecent droughts,mound densities of fire ant populationsalso diminished. One consequence of this reduction in fire ant numbershas beenthe difficulty in obtaining large S. irvicta colonies with appropriatecaste ratios and brood for sustaining/maintaininglaboratory cultures ofP. tricuspis. 32 Laredol Corumba2 ( Pantanal ) o ho F{ Cuiba2 l0) 24 .pl{ ( Pantanal ) da OrJ Ed ll ')n c0, 6Pl ntF E0) H Pres. Rogue Saenz Penaz (Chaco) 50 100 1-50 200 Precipitation (run) 1Data from NOAA (2001) ?n^L^'udvq f rom Schwerd,tf eqer (I97 6) FIG. 1. Climatograms of relatively harsh areasof Brazil (Pantanal) and Argentina (Chaco) occupied by solenopsis and Pseudacteon compared to Laredo, Texas, one of our pseudacteon releasesites. Average conditions are both hotter and drier at the Texas site. S. porter (pers. comm.) suggestedthese South American sites as possibly comparable to those in Texas. We are aware of no climatic zones in Argentina or Brazil with S. invicta and pseudacteon that can match the extremes of temperature and moisture (see for example, Fig. 1) and lack of predictable dry and wet periods characteristic of central and southern Tei*ar. Ho*e.rrer, the climatic pattems of some South American regions are more similar to those of Texas than are the source areas for P. nicuspis stocks currently being released in the US' Our workingassumption is thatphorid species or biotypesfrom temperateareas will be betterable to deal with Texasweather and climatethan can tropicalP. tricuspis from north of the
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