ECOLOGY OF LARRA BICOLOR (HYMENOPTERA: SPHECIDAE) IN THE
NORTHERN GULF
By Cheri Muthirakalayil Abraham
A Thesis Submitted to the Faculty of Mississippi State University in Partial Fulfillment of the Requirements for the Degree of Masters of Science in Entomology in the Department of Entomology and Plant Pathology
Mississippi State, Mississippi December 2008
Copyright by
Cheri Muthirakalayil Abraham
2008
ECOLOGY OF LARRA BICOLOR (HYMENOPTERA: SPHECIDAE) IN THE
NORTHERN GULF
By Cheri Muthirakalayil Abraham
Approved:
______David Wayne Held Gregg Munshaw Assisstant Professor of Entomology and Assisstant Professor of Plant and Soil Plant Pathology Science (Director of Thesis) (Minor Professor & Committee Member)
______M. Blake Layton Jr. Clarence Collison, Professor and Head, Extension Professor of Entomology and Department of Entomology and Plant Pathology Plant Pathology & Graduate Coordinator (Committee Member)
______William Kingery, Professor and Graduate Melissa J. Mixon, Coordinator, Plant and Soil Science Interim Dean of College of Agriculture and Life Sciences.
Name: Cheri Muthirakalayil Abraham Date of Degree: December 12, 2008 Institution: Mississippi State University Major Field: Turfgrass Entomology Major Professor: Dr. David W. Held Title of Study: ECOLOGY OF LARRA BICOLOR (HYMENOPTERA: SPHECIDAE) IN THE NORTHERN GULF Pages in Study: 62 Candidate for degree of Master of Science
Mole crickets (Scapteriscus spp.) are the most destructive pests in southern turf
and pasture grasses. In response to extensive losses from mole crickets, Florida formed a
task force to identify natural enemies in the native range of these pests. Two parasitic
insects, Larra bicolor and Ormia depleta, and Steinernema scapterisci, an entomopathogenic nematode, were imported and released. Of the two insects, only Larra bicolor has spread to other states infested with mole crickets in the southeast. The present study documents the seasonal biology of Larra bicolor in the northern Gulf region, ornamental plants that can be used as nectar sources, and the impact of these nectar sources on longevity of the wasp and parasitism of mole crickets. Results of field and laboratory experiments showed that the ornamenal Pentas lanceolata attracted wasps in the field and provided comparable or better longevity than Spermacoce verticillata which was the only known nectar source.
Key words: Larra bicolor, nectar sources, Mole cricket Bio control, Spermacoce
verticillata, Scapteriscus control.
DEDICATION
I would like to dedicate this document to my parents, M. C. Abraham and Elizabeth
Abraham, brother Tomci and fiancée, Tina.
ii
ACKNOWLEDGEMENTS
The author expresses his sincere gratitude to the many people without whose assistance this thesis would not be possible. First of all, I thank Dr. David W. Held, my graduate advisor, who worked endlessly to make this research possible. I will always be appreciative for his guidance and motivation which has been a never ending process.
Second, I would like to thank my committee members, namely, Dr. Gregg C.
Munshaw, Dr. Blake Layton Jr. and Dr. Clarence Collison for their unending support of this research and Dr. Dennis Rowe who helped me with part of the statistical analysis.
Special thanks are due to Corey Wheeler, research technician, for competent assistance in collecting data especially during times when I was on campus. Thanks to
Patrick Brousssard, Lloyd Easterly, and the staff of the Coastal Research and Extension
Center, Biloxi for making my stay there comfortable and keeping me at home with the facilities provided.
I would like to express my gratitude to the US Golf Association for funding this interesting project. The Green Section Research committee was encouraging and their comments helpful. Many thanks to the staff at Great Southern Golf Club, Gulf Hills Golf
Club and St. Andrews Golf Club where I frequented every week or two. I also thank the
Alabama Turfgrass Research Foundation for funding the survey for Larra bicolor in
Alabama. The staff at golf courses in Alabama namely Springhill College, Country Club
iii
of Mobile, Azalea City Golf Course, Timber Creek and Golf Creek Golf Club, Kiva
Dunes, Peninsula Golf Clubs and Golf Club at the Wharf were helpful during the survey.
Thanks to all of them.
The Mississippi Entomological Association was a strong source of
encouragement through the opportunities for paper presentations and scholarships, and
getting to know other students and accomplished professionals in the state and region. I
would like to thank Arlie and Ethel Wilson for the scholarship I received. Many thanks
to each and every one whose name I may have unknowingly omitted, but have been
instrumental in making my stay at Mississippi State University an experience which will be cherished throughout my life. Thank you one and all.
iv
TABLE OF CONTENTS
DEDICATION ...... ii ACKNOWLEDGEMENTS ...... iii LIST OF TABLES ...... vii LIST OF FIGURES ...... viii CHAPTER I. GENERAL INTRODUCTION ...... 1 Introduction to Turfgrass ...... 1 Turfgrass Pest Management ...... 4 Mole Crickets ...... 5 Pest Species, Distribution and Origin ...... 6 Damage by Mole Crickets ...... 8 Chemical Control ...... 9 Life Cycle and Biology of Scapteriscus vicinus and S. borellii ...... 10 Natural Enemies and Biological Controls...... 13 Larra bicolor- Mole Cricket Killers ...... 16 Morphology of Larra bicolor ...... 16 Parasitism and Biology of Larra bicolor ...... 17 Host Specificity of Larra bicolor ...... 19 Nectar Sources for Larra bicolor ...... 20
II. SEASONAL AND DIEL ACTIVITY OF LARRA BICOLOR IN THE SOUTHEASTERN UNITED STATES ...... 21 Abstract ...... 21 Introduction ...... 22 Materials and Methods ...... 22 Results ...... 25 Discussion ...... 30
III. ORNAMENTAL PLANTS AS NECTAR SOURCES FOR LARRA BICOLOR ..32 Abstract ...... 32 v
Introduction ...... 33 Materials and Methods ...... 37 Source of Plants and Wasps ...... 37 Evaluation of Flowering Plants in a Common Garden Experiment ...... 37 Evaluation of Wasp Longevity in the Laboratory ...... 39 Results ...... 41 Evaluation of Flowering Plants in a Common Garden Experiment ...... 41 Laboratory Longevity Assays ...... 42 Discussion ...... 43
IV. FIRST REPORT OF LARRA BICOLOR (HYMENOPTERA: SPHECIDAE) IN ALABAMA ...... 46 Abstract ...... 46 Introduction ...... 47 Materials and Methods ...... 49 Results and Discussion ...... 50
V. CONCLUSIONS...... 54 BIBLIOGRAPHY ...... 56
vi
LIST OF TABLES
Table Page
2.1 Locations Selected for Monthly Monitoring of Larra Wasps ...... 23
2.2 Diel Activity of Larra bicolor ...... 28 3.1 Horticultural Plants Evaluated as Nectar Sources for Larra bicolor ...... 38
4.1 Numbers of Larra bicolor Reported from Golf Courses in South Alabama ...... 51
vii
LIST OF FIGURES
Figure Page
1.1 Growth Pattern of Cool Season Turf (Turgeon 2005) ...... 2 1.2 Growth Pattern of Warm Season Turf (Turgeon 2005) ...... 2 1.3 Differences in Dactyls Spacing and Pronotum Coloration in Scapteriscus Mole Crickets ...... 8 1.4 a. Larra bicolor b. Larra analis (photo credit E. Trammel, AR) ...... 17 2.1 a. Larra bicolor Female b. Larra bicolor Male ...... 24 2.2 Seasonal Activity of Larra bicolor ...... 26
3.1 Larra bicolor Feeding on Pentas lanceolata ...... 35
3.2 Spermacoce verticillata as a Weed on the Roadsides of Florida ...... 36 3.3 The Garden Experiment with Plants in Bloom ...... 39
3.4 Arena for the Longevity Tests ...... 40 3.5 Field Test Results of Ornamental Plant Preference for Larra bicolor...... 41 3.6 Lab Trial I - Choice Test with No Food Control, Spermacoce and Pentas ...... 42 3.7 Lab Trial II with Spermacoce, Pentas and Sugar Water Control ...... 43
4.1 a. Larra bicolor (15–20 mm long) is Shown Here Feeding on Flowers of Spermacoce verticillata b. Stinging a Mole Cricket...... 48 4.2 a. A Female L. bicolor Preparing to Enter the Mound in Search for a Mole Cricket. b. Mole Cricket Mound with Hole that was Commonly Observed on Sites where L. bicolor Wasps were Active.. 52
viii
CHAPTER I
GENERAL INTRODUCTION
Introduction to Turfgrass
Ofth 7500sp ci sofplantsinfamilyPoac a (=Gramina ),onlyabout40 sp ci sar us dforth variousfunctionsofturf(Tashiro1987).Th commonlyus d t rm,‘turfgrass’r f rstoth plantofgrass,whil ‘turf’r pr s ntsacommunityof turfgrass sincludingth m diumofitsgrowth(Turg on2005).Accordingtoth climaticdiff r nc s,turfgrass sar mainlydivid dintowarms asongrass sandcool s asongrass s.Th ydiff rinth irr quir m ntofclimaticcondition,photosynth tic
ffici ncyandoth rgrowthr lat dproc ss s.Warms asongrass s(C4plants)r quir
27–35°C(80–95°F)foroptimumgrowth,wh r ascools asongrass s(C3plants)r quir
16–24°C(60–75°F)foroptimumtopgrowthand4–16°C(40–60°F)forrootgrowth
(Turg on2005).AlthoughC4plantsar consid r dmor photosynth tically ffici nt,C3 plantsshowlimit dornowint rdormancy(Turg on2005).Th diff r nc inth growth patt rnofth s twograsstyp sisshowninFigur 1.1and1.2.
1
Figur 1.1
GrowthPatt rnofCoolS asonTurf(Turg on2005)
Figur 1.2
GrowthPatt rnofWarmS asonTurf(Turg on2005)
2 Accordingtoclimat andpr cipitation,th contin ntalU.S.isdivid dintosix
turfgrasszon s(Pott r1998).Mississippiisinzon 3andcom sund rth warmhumid
zon .Ev nthoughth ntir stat isclassifi dund rzon 3,th climaticconditionsdo
varyalongth l ngthandbr adthofth stat .Th ar aalongth GulfCoast xp ri nc s
asubtropicalclimat whichiswarmandhumid,butplac snorthofth coast xp ri nc
cool rwint rsandwarm rsumm rs.Th tropicalclimat inth coastalr gionmak sit b stforth growthofwarms asongrass s.Bahiagrass(Paspalum notatum) , b rmudagrass(Cynodon spp.),zoysiagrass(Zoysia spp.),St.Augustin grass
(Stenotaphrum secundatum),andc ntip d grass(Eremochloa spp.)ar th maingrass s
growninthisr gion.Conditionsinth north rnpartofMississippiar mor favorabl
forzoysiagrass,wh r asconditionsinth south rnpartofth stat ar mor suit dto
growthofbahiagrassandSt.Augustin grass(Tashiro1987).C ntip d grasshas
distributionallov rth stat mostlyinhom lawnswhil b rmudagrassisus dingolf
cours sandoth rar as.
Unlik rowcropsandoth rplantsofcomm rcialimportanc ,turfgrassandits
importanc ar oft nov rlook d.Yi ldsandloss sinturfgrassar notasw usuallys
intraditionalcropsorcroppingsyst ms.Turfgrasshasmor ofana sth ticvalu andits b n fitsar notoft ntangibl .For xampl ,h r ar af wqualiti softurfwhichmak
th mimportant(Morris2003):
• Mod rat lysiz dlawnsof ighthom sar capabl ofcoolingair quival ntto70
tonsofairconditioning.
3 • A 232.3 m2 (2500 foot2) lawn absorbs carbon dioxid and produc s nough
oxyg ntosupportafourm mb rfamily.
• Ah althylawnr duc ssurfac wat rrunoffupto80tim s.
• Ab autifullawnincr as shom prop rtyvalu sby15%.
Apartfromth s statistics,turfgrassb autifi smillionsofhom lawns,isasaf playingsurfac onov r700,000athl ticfi lds,andprovid sasourc ofoutdoor
r cr ationforov r26milliongolf rsonabout17,000golfcours s.Witht nsof
thousandsofsodproduc rs,lawncar op ratorsandlandscap rs,th r ar atl ast20
millionha(50millionacr s)ofmanag dturf,withan stimat d$45billionindustryin
allformsinth U.S.(B ardandGr n1994).InMississippi,th r ar mor than1
millionha(2.5millionacr s)ofmaintain dturf,withatl ast121,405ha(300,000acr s)
oflawnsurrounding740,000hom s,160golfcours sand2,000athl ticfi lds
(Anonymous2005).Forthisr ason,turfgrassisan conomically@importantindustry.
Turfgrass Pest Management
Accordingtoth functionsturfp rforms,th tol ranc l v ltop stsanddis as s
vary.Th tol ranc top stsonlowbudg tgolfcours s(i. .,municipalorpublicgolf
cours s)wouldb mor thanthatofprivat ors mi@privat golfcours s.Similarly,th
tol ranc l v lforp stsinhom lawnsdiff rfromthatofanathl ticfi ld.Privat or
s mi@privat golfcours swouldhav littl ornotol ranc forp stsandtypicallyus
mor ch micalstocontrolp ststhanpublicgolfcours s.
4 P stsanddis as svaryaccordingtoth climaticzon inwhichth turfisgrown.
For xampl ,mol crick tsar commonlyagr duponasth mostm nacingp stin
turfgrassinth south rnorsouth ast rnpartsofth U.S.,butofnoconc rninth
north rnU.S.wh r whit grubsar mor probl matic.Apartfromf rtiliz rs,annual
xp nditur sonturfgrassar mostlyforplantprot ctionch micalssuchash rbicid s,
fungicid s,andins cticid s. D spit an mphasisonint grat dp stmanag m nt,turf
manag rsmayb mor lik lytochoos ach micalcontrolthanrisklosingafairwayor puttinggr n.As vid nc ofthis,125,000haofhighmaint nanc ar asofgolfcours s
lik t s,gr ns,andfairwaysw r spray dwithins cticid sin1996(Rack 2000).By
comparison,31%ofp sticid sus dbylawncar compani sar ins cticid sv rsusonly
19%ongolfcours s(GCSAA1998,Rack 2000).Th targ tsofth s ins cticid s,
particularlyinth warmhumidr gionar p stmol crick ts(Scapteriscus spp.),chinch
bugs(Blissusspp.),andcat rpillarp stslik sodw bworms,armyworms,cutworms,and
fir ants.
Mole Crickets
“Mole crickets are the most destructive insect pests of golf courses, lawns and sod
farms in the southeastern United States. These pests account for hundreds of millions of
dollars in damage and control costs every year” (Pott r1998).Ext nsiv ar asplant din
bahiagrassandb rmudagrass,accompani dbysandysoilswhichaidandfavorth ir
tunn lingb havior(Tashiro1987),mak th GulfandAtlanticcoastsb stsuit dformol
crick ts.
5 Pest Species, Distribution and Origin
Ofth 80sp ci sofmol crick tsworldwid ,t noccurincontin ntalU.S.,
Hawaii,Pu rtoRico,andVirginIslandsnam ly;Neocurtilla hexadactyla(P rty)th north rnmol crick t,Gryllotalpa cultrigerUhl rth w st rnmol crick t,Gryllotalpa majorSaussur th prairi mol crick t,Gryllotalpa gryllotalpa(L.)Europ anmol crick t,Gryllotalpa orientalisBurm ist rori ntalmol crick t,Scapteriscus abbreviatus
Scudd rshortwing dmol crick t,Scapteriscus borelliiGiglio@Tos(=S.acletus)south rn mol crick t,Scapteriscus didactylus(Latr ill )W stIndianmol crick t,orchanga,
Scapteriscus imitatusNickl andCastn rimitatormol crick t,andScapteriscus vicinus
Scudd rtawnymol crick t(Nickl 1992).Amongth s ,onlyfoursp ci sar common inth south ast rnU.S.;th north rnmol crick t,anativ sp ci s,andth immigrant sp ci s; Scapteriscus vicinus,S. borellii,andS. abbreviatus.Th north rnmol crick tis afourclaw dmol crick tunlik th oth rthr whichar twoclaw d(Frank tal.
1994).Th north rnmol crick tison of35sp ci soffourclaw dmol crick ts occurringinthr g n raNeocurtilla, Gryllotalpella, Gryllotalpa (Nickl andCastn r
1984).
Short@wing dandsouth rnmol crick ts,w r introduc dintoBrunswick,
G orgiain1904,wh r astawnymol crick twasintroduc dhalfad cad arli ratth sam plac (Walk randNickl 1981).Th yar b li v dtohav immigrat dtoth U.S. byhitchhikinginth ballastsofth ships,aft rwhichth yslowlyspr ad.Th r ar two morphologicallyr cognizabl formsofth south rnmol crick t,four@dotformand mottl dpronotumform,whichw r introduc ds parat ly.Th formwithamottl d
6 pronotum nt r dth U.S.throughSouthCarolinain1916,T xasin1926,andFloridain
1920(Nickl andCastn r1984).Sinc th irwingsar v stigial,short@wing dmol
crick tscannotnaturallyspr adfarandar mostlyprobl maticinsouth rnFlorida.In
Mississippi,th south rnandtawnymol crick tsoccurinth south rnpartofth stat
typicallysouthofI@20(H ld,p rsonalobs rvation).
Dactylspacingisamaincharact rwhichh lpsdistinguishb tw nS. vicinus and
S. borellii.Th twotibialdactylsonS. borellii ar s parat dbyalmostth sam widthas
ith rofth twoclaws,wh r asth tibialclawsonS. vicinusar s parat dbyasmall r
distanc givinganappar ntV@shap toth tawnymol crick tandaU@shap toth
south rnmol crick t(Hayslip1943)(Figur 1.3).
7 Figur 1.3
Diff r nc sinDactylsSpacingandPronotumColorationinScapteriscusMol Crick ts (photocr dit,L.Buss,Univ.ofFL).
Damage by Mole Crickets
Withth irstrongfrontl gs(fossorial),mol crick tscaus thr typ sofcaviti s inth ground.Tunn lsr f rtoth d p rmin sth ymak inth ground.Gall ri sar th horizontalmin smad justb lowth soilsurfac whichcaus soiltobulg abov th surfac .Boths x scanmak tunn lsandgall ri s,but ggchamb rsar sp cialty caviti smad bygravidf mal s.Th yoccuratd pthsvaryingfrom10@30cmsb lowth
8 soilsurfac .So,inth soilprofil b ginningatth surfac ,w would ncount rgall ri s first,th ntunn ls,andlat r ggchamb rs.Gall ri sar damagingingolfcours putting gr nsb caus th yd fl ctth rollofagolfball.H avyrainsandirrigationcancl ar th s gall ri sinsandysoilssofr shdamag isoft nappar nt.
Apartfromth tunn ling,h rbivorycanalsocaus damag (Frank tal.1994).
Gut xaminationofth south rnmol crick tsshowthatth irdi tmostly(91%) constitut sofins cts,wh r astawnymol crick tshav aplant@richdi t(66%)with ins ctsoranimalsconstitutingasmall rportion(Taylor1979).Oth rScapteriscusmol crick tsar similarinf dinghabitsandmainlyfallintoth s twocat gori s(Castn r andFowl r1984).
Rootdisruptionthroughtunn lingcaus sdryingofth soilanduprooting follow dbyd siccationofinf st dgrass.Exc ssiv tunn lingcanw ak norkillastand ofturfgrass(Adj i tal.2003).Ev nthoughsp ci smaydiff rbyth charact ristic damag th ymak ,allsp ci sdocaus m chanicaldamag (Castn randFowl r1984).
Incr as ddamag isacc ntuat dinmoist(H rtlandBrand nburg2002)andsandysoils.
Asar sult,w ll@wat r dturfgrass, sp ciallyongolfcours s,ar mor vuln rabl to mol crick tdamag (Hayslip1943).
Chemical Control
B ingsubt rran anp sts,mol crick tsar not asytocontrol.Tob ff ctiv , ch micalsmustp n trat th thatchlay randr achth mol crick tsinados which wouldb l thaltoth m.Thismayr quir multipl applicationsofp sticid stok p
9 th mund rch ck.Mol crick tscanr port dlyd t ctandavoidsoilstr at dwith ins cticid s(Brand nburg2002)furth rconfoundingmanag m nt.W canobs rv larg amountsofdamag duringth spring,th r for ch micalapplicationsar n c ssaryat thattim ,how v rth fficacyofp sticid sar imp d dbyth larg nymphsandadults abundantlypr s ntatthattim .Sinc th ins cticid wouldb mor ff ctiv against young rnymphs,th yar tob targ t dforoptimumr sults.Thisgiv sawindowfrom
Jun throughJulyforapplicationofth p sticid s,b caus nymphsar smalland asily controll dbutdamag isnotobvious.Thisp riodhow v r,isth b stchanc tocontrol mol crick ts(H ldandLayton2006).Baitsar ins cticid sformulat donapalatabl foodformol crick ts.Wh nappli dinAugust,baitsw r foundtob mor ff ctiv thanoth rmod sofapplication(ShortandKo hl r1977).Amongcommonins cticid s, fipronil(ChipcoChoic ,Bay rEnvironm ntalSci nc )isth most ff ctiv .
Life Cycle and Biology of Scapteriscus vicinus and S. borellii
Mol crick ts ggsar dirtybrownorgrayish,whichar initiallywhit ,ovaland unsculptur d.Inth ggchamb r,f mal slayaclutchof ggs,av rag numb rof35 p rclutch.Onc ggsar laid,f mal susuallydonotr visitth chamb r,unlik
Neocurtillia hexadactylaf mal s,whichdor visitth chamb r.Inc ntralFlorida, gg layingforbothsp ci sb ginsbyth latt rhalfofMarchandismostlycompl t dbyth firstofJuly(Hayslip1943).Ev ryactivityofth mol crick tsfromflightp riodto ovipositionand gghatchwillshowdiff r nc sdu tog ographiclocation,andh nc maynotb th sam inplac swithdiff r ntlatitud s.Incubationof ggsist mp ratur
10 d p nd nt.Incubationisshort stwh nsoilt mp ratur sar around29°C(84°F)and
long stwh nsoilt mp ratur saround20°C(69°F).Incubationistypically5dshort r
forS. vicinus(Hayslip1943).
Eggshatchintonymphs,whichar similarinapp aranc toadults, xc ptth y
lackwings.Inc ntralFlorida,hatchofS. vicinusoccursduringlat April,whil ggsof
S. borellii hatchin arlyMay(Hayslip1943).Onc m rg d,nymphsb gins archingfor
food,f dingon ggs,chorions,and v nw ak rnymphs.Nymphsmolt6–10tim s b for b comingadults.Th diff r ntmoltscanb r cogniz dbypronotall ngthwhich
rang sfrom[email protected]mm(0.07–0.32in)(Hudson1987).Numb rofmoltsandnumb r
ofdaysr quir dforb comingadultsvari sb tw nsp ci s.Th d v lopm ntof
wingpadsisanoth rcharact rassociat dwithth d v lopm ntstag sinmol crick t.
Th thr stag sofwingpadd v lopm ntar abs nc ofwingpads,pr s nc ofsmall
wingpads,pr s nc oflarg wingpads.On ortwoofth p nultimat nymphalstag s
willhav smallwingpadswh r asth lastnymphstag haslarg wingpads(Hudson
1987).South rnandtawnymol crick tsar univoltin .How v r,aportionofth populationmayb com adultsb for wint r,wh r asmostov rwint rasnymphs
(Hayslip1943).
Cool rsoilt mp ratur scaus th mtor mainund rgrounduntilfavorabl
conditionsallowth mtostartth irnuptialflights.Inamarkandr captur study,on
mol crick twascoll ct d8kmsfromth r l as sit ,althoughth yar nottypically
longdistanc fli rs(Frank tal.1994).Adultmol crick tshav twopairsofwings
whichar cov r dbyan tworkofv ins,whichar toughtub ssupportingth wing
11 m mbran s.Spac sb tw nth v insar call dc lls.For wingsofmal sandf mal s
diff rslightlyinthatfor wingsofmal shav apairoflarg c lls.Th ant riorsid sof
th s c llsar harp@shap dandcall dth stridulatoryar a.F mal slacksuchlarg c lls.
Inmal s,on ofth v inson achwingismodifi dwithalin oftinyt thtoforma
stridulatoryfil .Thisfil isdrawnacrossascrap ronth oth rwing.Asth wings
clos ,thismak sasound,whichisth irmatingsong(i. .stridulation).Mal sr p atthis
songbyop ningandclosingth irwingsinrapidsucc ssion.Th arrang m ntofth
t thinth stridulatoryfil diff rsfromsp ci stosp ci s,soth songdiff rsfrom
sp ci stosp ci s.Th loudn ssofth songsvari sbutistypicallyabout70dBat15cm
fromth sourc .Th songisamplifi dbywid ningth mouthofth irgall ryintoa
funn lshap .Th h aringorgans(tympanum)ofmol crick tsar onth tibia ofth
for l gs(Frank tal.1994).
Mal tawnyandsouth rnmol crick tssingatnighttoattractf mal s.Flights
b ginsoonaft rsuns tand ndaft rabouton hourinbothth tawnymol crick tand
south rnmol crick t(Hayslip1943,Ulagaraj1975).Th r ar twoflightp riodsfor bothsouth rnandtawnymol crick ts;amajorspringflightandminorfallflightbothof
whichoccurwh nhight mp ratur sar ov r18°C(Ulagaraj1975).Infall,how v r, south rnmol crick tsar pr dominantlyfound(Hayslip1943).Th p akoftawnymol crick tflightisMarchto arlyApril,andSouth rnmol crick tflightisAprilto arly
May(Hudson2001).About14daft rmating,th f mal lays ggs(Hudson2001).
12
Natural Enemies and Biological Controls
Inth nativ rang ,Scapteriscus mol crick tsar uncommonandrar lyofp st status(Hudson tal.1988).Thisislik lydu toth pr s nc ofnatural n mi swhich k pth irpopulationsinch ck.Similarly,c rtainpr datorshav b nnot dfor attackingmol crick tsinth Unit dStat s.Subt rran anpr datorslik groundb tl s fromth g nusPasimachusandassassinbugsofth g nusSirthenea attackadultsand nymphs.Adultsofbombardi rb tl sp ci s,Stenaptinus jessoensis (Morawitz)ar sp cialistpr datorsofmol crick t ggs(Hudson tal.1988).Oth rgrounddw lling pr datoryins ctsandspid rsknowntoattackmol crick tsinclud arwigs
(D rmapt ra),groundb tl s(Carabida ),tig rb tl s(Cicind llida )andspid rsinth famili sLycosida andSalticida (Hudson tal.1988).TwoS.vicinuscaptur din acoustictrapsinN wOrl answ r foundtob parasitiz dbyAcridomyiaspp.(Dipt ra:
Anthomyiida )whichar usuallyparasitoidsofgrasshopp rs(Acridida )(H nn and
Johnson2001).Birdslik sandhillcran sandcattl gr tsforag onmol crick tswh n
th yar n artoth surfac .Insom instanc s,foragingofins ctivorousanimalslik
armadillosdomor damag toturfthanth mol crick tsth ms lv s.Amongoth r
v rt brat pr datorsar fox s,skunks,raccoons,toads,andbirds(Sailor1984)which
awaitth mabov ground.
Apartfromth s pr dators,mol crick tpopulationsar subj ct dtoattackby
pathog nslik bact ria,fungi,virusandprotozoans.Fungalpathog nslik Beauveria
bassiana (Balasamo),Isaria spp.,Metarhizium anisopliae(M tchnikoff),Entomophthora
13 spp.,andSorosporella uvellahav b nfoundtoinf ctmol crick ts(Hudson tal.
1988). Entomopathog nicn matod Steinernema (=Neoaplectana)carpocapsae W is r whichattacksawid rang ofins ctsw r alsofoundtoinf ctmol crick ts(LimaCosta
tal.1984).
D spit thislistof nd micornaturaliz darthropodsandpathog nswhichattack
Scapteriscus mol crick ts,th irpopulationsw r n v rsignificantlyr duc d.In
Florida,amol crick ttaskforc wasform dtoinv stigat classicalbiologicalcontrolof mol crick tsusingnatural n mi sfromth irnativ rang .Th classicalbiocontrol ag ntswhichhav b nsucc ssfullyintroduc dand stablish dinFloridaar th r d@
y dflyOrmia depleta(Dipt ra:Tachinida ), b n ficialn matod Steinernema scapterisci, andth waspLarra bicolor(Sph cida ).InFlorida,acoustictrapcaptur sof
Scapteriscus mol crick tsov r25y arsonth sam sit sb for andaft rth s biologicalcontrolsw r stablish dindicat a99%r ductionintrapcaptur s(Frankand
Walk r2006).Thishasb nattribut dlarg lytoth succ ssofth introductionofth s biologicalcontrols.
InJun 1939,Scapteriscus mol crick tscoll ct dfromB l m,Brazilw r found tob parasitiz dbyatachinidfly(Wolcott1940).In1983,gravidf mal softh sam sp ci sr spond dtosynth siz dcallsofScapteriscus borelliiinParaguay(Fowl rand
Kochalka1985) arli rr port d(Mangold1978).Thisflywasfirstid ntifi das
Euphasiopteryx australis (Wolcott1940)andlat rfoundtob Euphasiopteryx depleta
(Sabrowsky1953).Th g n ricnam waslat rmad asynonymofOrmia, thus changingth nam toOrmia depleta(Wood1987).Aft rcoll ctingth flyandhosts
14 fromPiracicaba,Brazil,th firstlaboratoryr ar dcolonywasachi v din1987.In spring1988,adultOrmia depletaw r r l as dinFlorida(Frank tal.1996).Th s fli s hav b nsucc ssfullyr ar donallsp ci sofScapteriscusmol crick tsandontwo gryllids(AnurogryllusandGryllus)(Fowl randM sa1987).Ext nsiv r s archand r l as softh tachinidflyinFloridasucc d din stablishinglocalpopulations.Injust on y ar,th r wasan85%d cr as inmol crick tinf stationscompar dtoar as wh r th flyhadnotb com stablish d(Frank tal.1996).Byth summ rof1994,O. depletahadspr adthroughoutFloridaandthisspr adwasassociat dwithd cr as sin mol crick tpopulations(Frank tal.1996).AcoustictrapsinN wOrl ans,LAhav trapp dS. borellii whichw r parasitiz datagr at rrat thanwas arli rr port dfrom
Florida(Frank tal.1996,H nn andJohnson2001).
In1985,Steinernema scapterisci wasbroughtfromUruguaytoFlorida.Aft r laboratory valuation,th firstsmallplotr l as sw r don n arth Univ rsityof
FloridainAlachuaCountyinJun 1985(Parkman tal.1993).Th r ar thr s parat lif stag s: gg,fourstag sofjuv nil s,andadult(mal andf mal )ofwhichth third stag juv nil isth mostimportantforbiologicalcontrolofmol crick ts.Th y nt r th mol crick tthroughitsmouthorspiracl s.Onc insid ,th juv nil sr l as bact riaintoth h moc lofth irhost.Th bact riumcaus sd athofth mol crick t bymultiplyingwithin,andth n matod f dsonth bact ria(Nguy nandSmart1992).
Initiallyr l as dasaclassicalbiocontrolag ntorbiop sticid ,thisn matod hasb n comm rcializ dandnowsoldund rth trad nam N matacS™.Th succ ssful inoculativ r l as ofthis xotic ntomopathog nicn matod wasth firstclassical
15 biocontrolagainstmol crick ts(Parkman tal1994,ParkmanandSmart1996).Ev n thoughth initialcostsofusingN matacSisconsid rablyhigh rthanth costofan ins cticid tr atm nt,inf ctionmayp rsistfromy artoy ar.Ev nthoughstudi shav shownthatmol crick tscanb haviorallyavoidsoilstr at dwith ntomopathog nic fungi(Villani tal.2002),aft rSteinernema w r stablish d,upto80%ofmol crick tstrapp donthos sit sw r inf ct d(L ppla tal.2007).Thisn matod isalso compatibl withins cticid scommonlyus dinturfgrass(BarbaraandBuss2006),
Larra bicolor- Mole Cricket Killers
Ov r60sp ci softh g nusLarra(Hym nopt ra:Sph cida )ar knowntob parasitoidsthatpr yonmol crick ts(BohartandM nk 1976).Larra americana
Saussur nowknownasLarra bicolor(BohartandM nk 1976)wasintroduc dinto
Floridathroughth ffortsofth mol crick ttaskforc .Succ ssfulintroduction
(Wolcott1938)and stablishm nt(Wolcott1941)of Larra bicolor intoPu rtoRico trigg r dth s archforthisnatural n mytocontrolth damag caus dbyp stmol crick ts.
Morphology of Larra bicolor
Th abdom nofLarra bicolorissolidr dandwingsar smokybrownorindigo blu withsilv rywhit markingsonth irh ad.Incontrast,Larra analis,anativ sp ci s thatattacksNeocurtilla hexadactyla, hassimilarcolor dwingsandadarkcolor d abdom nwithonlyr datth tip(Figur 1.4).
16 Th tarsalclawsar shortandwithoutinn rt th.Th for tibiahasspin rows, withoutt thoninn rmarginofmandibl andpropod umisconstrict datth l v lof spiracl swithaglossyint gum nt.F mal shav athick rpronotalcollar,straightontop andashighasscutumandst rnumIIwithoutapairofsp cialflatt n dovalar as basally.Th pygidialplat isbar andwithoutatransv rs rowofshorts ta apically.
F mal sar g n rallybigg r(av rag siz 22mm)thanth irmal count rparts(Frank andSourakov2002).
Figur 1.4
a.Larra bicolor b.Larra analis(photocr ditE.Tramm l,AR)
Parasitism and Biology of Larra bicolor
Inshort rmownturfgrass,f mal Larrawaspsmak shortflightsacrossth grass landingth ninitiatingawalkings archforhosts(p rsonalobs rvations).Mosthost s archingb haviorsobs rv dinMississippiw r oncloudl ssdayswithlowtomod rat winds(i. ,≤10mil p rhour),whichisconsist ntwithWolcott(1938).Also,Larraar
17 g n rallyfirstobs rv dinMississippiin arlytomidmorningcontinuingthroughth lat aft rnoonifconditionsar favorabl (p rsonalobs rvations).Th r isvariation r port dinth foragingb haviorofLarrawaspsobs rv dbydiff r ntr s arch rsin diff r ntlocations.InBrazil,flightabundanc wasr lativ toth b ginningand ndof th rainys ason(F bruaryandMay,r sp ctiv ly).Thisisattribut dtoth as of locatinghosttunn lsinth moistsoilduringth s tim s.Soilandambi ntair t mp ratur smayalsoplayarol inhuntingb havior.Assoilt mp ratur sincr as dto
50°C,hosthuntingb cam sporadic(Castn randFowl r1987).Wh nwaspsar not huntingth ymays ksh lt rinth foliag ofn ctarproducingplantsorb haviormay switchfromhuntington ctarf ding.
Wh nhunting,Larrawaspswalkonth surfac ofsoilant nnatingth soiland turfasth ymov (p rsonalobs rvations).Wh nasurfac tunn lis ncount r dth waspwill nt rand xitth sam tunn lmanytim saft rwhich,du tounknown r asons,th mol crick trush soutofth tunn l,follow dbyth wasp(Castn rand
Fowl r1987).Althoughattacksmayoccurund rground,th waspismor fr qu ntly obs rv dattackingpr yonth soilsurfac .Itisr port dthatn arlyhalfofpr ythat xit th hol scap parasitism(Castn randFowl r1987).Th waspstingsth mol crick t andlaysan ggonc th pr yisinhand.Thiswaspisan ctoparasitoid,anditsv nom onlyt mporarilyparalys sth mol crick tprovidingawindowforoviposition.Aft r layingth gg,th waspmayl av th mol crick t(FrankandSourakov2002)or r maintoguardituntilth hostr cup rat sfromparalysisandr sum sitsnormalroutin
(Castn r1988).Instudi s,itwasobs rv dthatmajorityofth waspslaidonlyon gg
18 onth v ntralsid ofth mol crick t.Ifan ggwas arli rlaid,itwouldb r mov d b for anoth rwaslaid.Thisb havior nhanc dth chanc ofth singl larva succ ssfullyd v loping(Castn r1986).
Eggincubationmaytak aslittl as4daysinsumm r,andth r ar 5larval instarsb for pupation.Tim fromovipositiontopupationist mp ratur d p nd ntand tak sfrom12–30days(Cushman1935,FrankandSourakov2002).About2w ksaft r oviposition,th wasplarva now14–26mm,willkillth host,consumingth softbody partsl avingonlyth scl rotiz dportions.Th cocoonisform dinaboutadayandis construct dofasilk ncas andcommonlycov r dwithsand.Th adultwasp m rg s
6–8w ksaft rcocoonsar form d(Cushman1935,FrankandSourakov2002).
Host Specificity of Larra bicolor
Larra bicolor willsucc ssfullyd v loponallsp ci sofScapteriscusmol crick tspr s ntinth Unit dStat sbutnotonth nativ sp ci s,N. hexadactyla.Th nativ mol crick twh nattack d mitsastickyanals cr tionwhich ntangl sth wasp, notallowingfurth rparasitism.Ev nwh nL. bicolor canr adilyovipositonN. hexadactyla,th wasplarvacannotsucc ssfullyd v loponthathost.Larra analis,th afor m ntion dnativ sp ci s,isth commonparasitoidofN. hexadactyla(Frankand
Sourakov2002).Havingco xist dsinc arly1900s,th nativ sp ci s,Larra analis hav noty tb nconsid r daparasitoidofScapteriscusmol crick ts.
19
Nectar Sources for Larra bicolor
Larra bicolor isr port dtof donflow rsofs v ralsp ci sofunr lat dw ds andwildflow rsincludingdollarw dandragw d(Frank tal.1994).Ar vi wof lit ratur showsth r hav b nnostudi stoid ntifyornam ntalplantswithflow rsthat canb us dbyL. bicolor. Amongth w dysp ci s,Spermacoce verticillata
(Rubiac a )ispr f rr dbyth waspandiscommonlyus dtomonitorwaspactivity
(M agh randFrank1998).Floralcharact risticsimportantforL. bicolorar floral d pth,whichis qualtoth l ngthofth irglossaincas ofSpermacoce(Ar valoand
Frank2005),andan ctarwithasucros :h xos ratio>1(Ar valoandFrank, unpublish ddata).
PopulationsofL. bicoloronflow rsar mal bias dinth morning,wh n pr sumablyf mal sar hunting,butcontainmor f mal sinth aft rnoonand arly
v ning(Castn randFowl r1987).Flow rf dingorforagingoccursfrom0830to
1530hoursformal andf mal L. bicolor(Castn randFowl r1987)butisd p nd nton t mp ratur .Inadditiontof ding,th s waspsmayalsor stonflow ringplants.Th originsorpurpos ofsuchb haviorsar notknown.
20
CHAPTER II
SEASONAL AND DIEL ACTIVITY OF LARRA BICOLOR IN THE
SOUTHEASTERN UNITED STATES
Abstract
Ev nthoughLarra bicolorhasb ncontrollingmol crick tsinFloridasinc
1993,th r hasb nnostudytodocum ntth s asonalanddi lactivityofth waspin
NorthAm rica.Th onlypr viousstudy(Castn randFowl r1987)thatdocum nt dth
di lactivitywasconduct dinPu rtoRicooritsnativ rang andh nc mayb diff r nt
fromwaspb haviorsinNorthAm rica.Th obj ctiv softhisstudyw r tor cordth
s asonalactivityofth waspinth north rnGulfCoastr gionandtor cordth di l
activityofforagingwasps.Ar asinsouthMississippiwh r populationsofLarra bicolor
w r knowntob stablish dw r surv y d v rytwow ksduring2006@2008to
docum ntth s asonalactivity.Inon ofth s ar as,hourlyobs rvationsw r don to
not th di lactivityofth wasp.Waspsw r typicallyactiv fromlat Mayto arly
D c mb rwithslightvariationsb tw nlocations.Waspsw r activ as arlyas0600h
andr main dactiv until1900h.Thisinformationisus fultoincorporat anint grat d
approachtocontrolmol crick tsbyr ducingins cticid sandtimingspraysinsucha
waythatwaspsar notaff ct d.
21 Introduction
Mostofth pr @r l as r s arch,includingbiologicaland cologicalstudi son
Larrawasps,wasconduct dinBrazilandPu rtoRico.Th mostwid lycit dstudyon th s asonalflightp riodanddi lpatt rnsofL. bicolor wasconduct dinPu rtoRico.
Thisstudy,conduct dinon g ographiclocationov r10dinasingl growings ason, do snotprovid informationaboutth activityofthiswaspinsubtropicalpartsofNorth
Am rica.Asth wasp xpandsfurth rnorthandw stinth south rnU.S.(H ld2005,
Abraham tal.2008),itisimportanttound rstanditsflightactivityinthisr gion.Th obj ctiv ofthisstudyh nc wastomonitorth s asonalactivityanddi lactivityof
Larra bicolorinth North rnGulfr gion.
MaterialsandMethods
FromJun –August2006, ightgolfcours sinsouthMississippiw r visit dand surv y dforL. bicolor.Manyofth s cours sinclud sit sinsouthMississippiwh r
Larrawasoriginallydiscov r d(H ld2005).Ofth s ight,thr (Tabl 2.1)w r s l ct dfors asonalmonitoringofLarra bicolor.Onth s thr cours s,twosit s wh r L. bicolor w r foundonmor thanon occasionw r s l ct dformonthly monitoring.
22 Tabl 2.1
LocationsS l ct dforMonthlyMonitoringofLarraWasps
Name, Location Site 1 Site 2
Gulf Hills Golf Club, 5th fairway 7th Tee Ocean Springs, MS St. Andrews Golf Club, 5th fairway 7th green Ocean Springs, MS Great Southern Golf 11th fairway Back 9 Club, Gulfport, MS MS State University, Flower garden Flower garden Coastal REC, Biloxi, MS
FromApril–D c mb r,2006–2008, achsit wasscout dfor15–20minut sfor waspactivity.On achgolfcours sit ,ar asinf st dwithmol crick tsandshowing fr shactivityw r firstscout d.Th s ar asw r s l ct db caus L. bicolor commonly
walksov rth turfs archingforactiv burrowsinwhichto nt r.Obs rvationsdon in
th gard natth CoastalRECw r ofwaspsforagingonflow ringplants(i. .,
Spermacoce verticillata orPentas lanceolata).Sinc th s waspsar not dn ctar
f d rs,monitoringflow ringplantswouldgaug th activityofL. bicolorinthat
location.Monitoringwasdon oncl ardays,b tw n1000–1500hours.Obs rvations
w r don v rytwow ks.
Di lactivityofL. bicolorwasstudi dbymonitoringwaspactivityonflow rsof
bloomingplotsofSpermacoceverticillata andPentaslanceolata.Th s plotsw r part
ofth gard n xp rim ntd scrib dinmor d tailinChapt rIII.Thisstudywas
conduct dfrom7–10Aug,2007and9@11Oct,2008wh nwasppopulationsw r
23 abundant noughtocoll ctdata.Th numb rands xofwaspsonflow ringplantsin
ightplotsinth gard nw r tak nhourlyfrom0500–2000h achday.F mal andmal waspsw r distinguish dbas donmorphologicaldiff r nc softh irwings.Th f mal
Larrawasphadasolidblackcolortoitswingandadarkr dabdom noppos dtoth lightr dabdom nandwinglight ratth bas anddark ratth ap xinth mal s(Figur
2.1)(p rsonalobs rvations).
Figur 2.1
a.Larrabicolor F mal b.Larra bicolor Mal
Duringth s xp rim nts,obs rvationsw r mad for20minut s achhour.
Waspsthatland donflow rsw r count d v nifth yd part dwithoutactually f ding.Dataw r coll ct dalldayd spit chang sinw ath rsuchasrainorov rcast conditions.T mp ratur datawastak nfromdatalogg rs(#450,Sp ctrum
T chnologi s,Plainfi ld,IL)adjac nttoth flow rgard n.
24 Results
Th s asonalactivityofLarra bicolorhasb nmonitor dsinc 2006andisinits thirdy ar.Typically,waspsw r firstobs rv dinMayandactivitycontinu duntil
D c mb r(Figur 2.2)wh nth ambi ntt mp ratur stypicallyw r atorb low20°C.
Waspsw r notobs rv dongolfcours sbrowsingonth turforparasitizingmol crick tsb for Jun oraft rNov mb r.Inth flow rgard n,how v r,waspsw r pr s nt arli randlat rinth s ason.Waspsw r activ onallsit ssurv y d xc pt on .Waspsw r pr s ntatSt.Andr wsgolfcours in2006butn v robs rv d th r aft r.
25
Figur 2.2
S asonalActivityofLarra bicolor
26 Waspsw r obs rv donflow rsfromabout0600hto1900h achdayin summ randfrom0900hto1700hinfallwh nt mp ratur sw r cool r(Tabl 2.2).
Sunris andsuns tduringsumm rdaysw r at0520hrsand1845hrs,and0600hrsand
1730hrsduringfall,r sp ctiv ly.Mal sw r consist ntlypr s ntat achhourly obs rvationwh r asf mal foragingwasmor sporadic.P akforagingonflow rsis hardtod t rmin b caus som waspscouldr mainonflow rsandnotforag .Wasps w r stillcount difpr s ntonth flow rsfromth pr viousobs rvation.Th gr at st numb rofmal swasr cord d(Tabl 2.2)at1800,1000,and1000–1100hoursforAug
7–9,r sp ctiv ly,wh r asth gr at stnumb rofmal sduringOct9@11w r at1500 hrsonallthr days.Th gr at stnumb roff mal swasr cord dat1100,1300,and
1000forAug.7–9,r sp ctiv lyandduringOct9@11abundanc off mal swasalso aroundth 1500hrmark.Waspsw r obs rv dr stinginth shad dint riorof flow ringplantsandjustaf ww r obs rv dtosp ndth ov rnightp riodonth plant.
Int r stingly,mostwaspsl av th plantatorn ardusk.Att mptstofollowindividual waspsatth ndofth dayw r notsucc ssful.Waspscouldb “Sl ping”in surroundingv g tationinaclust roralon ass ninoth rsolitarywasps(O'N ill
2001).
27 Tabl 2.2
Di lActivityofLarra bicolor
Numb rofwasps Dat Tim Mal F mal Total Soil Air T mp ratur T mp ratur (°C) (°C) 08@07@07 0500 0 0 0 28.7 27.6 0600 0 0 0 28.7 27.2 0700 1 6 7 28.3 27.2 0800 1 0 1 28.3 28.3 0900 7 0 7 28.3 30.4 1000 14 0 14 28.3 31.6 1100 15 0 15 28.7 34.1 1200 11 3 14 28.7 34.9 1300 16 14 30 29.5 34.9 1400 12 6 18 29.9 31.2 1500 1 4 5 28.7 24.4 1600 5 9 14 28.0 26.0 1700 25 5 30 28.0 29.5 1800 31 9 40 28.3 31.2 1900 15 5 20 28.3 30.4 2000 0 0 0 28.3 29.5 08@08@07 0500 0 0 0 28.0 27.6 0600 0 0 0 27.6 26.8 0700 14 6 20 27.6 27.2 0800 7 0 7 27.6 28.3 0900 13 2 15 27.6 29.9 1000 21 7 28 28.0 31.6 1100 7 1 8 28.0 34.1 1200 17 1 18 28.7 35.8 1300 18 10 28 29.5 33.6 1400 9 7 16 29.5 34.1 1500 8 2 10 29.9 34.9 1600 3 0 3 30.4 33.6 1700 2 0 2 30.4 33.6 1800 2 0 2 30.4 32.8 1900 4 0 4 29.9 32.4 2000 0 0 0 29.9 32.0 08@09@07 0500 0 0 0 28.7 28.0 0600 0 0 0 28.3 27.6 0700 0 0 0 28.3 27.6 28 Tabl 2.2Contd… 0800 0 0 0 28.3 29.5 0900 9 0 9 28.3 31.6 1000 18 11 29 28.7 32.8 1100 18 5 23 29.1 34.9 1200 4 1 5 29.5 34.9 1300 6 1 7 29.9 34.5 1400 3 5 8 30.4 34.9 1500 1 0 1 30.4 29.1 1600 4 0 4 30.4 31.6 1700 0 0 0 29.9 29.1 1800 1 0 1 29.5 27.6 1900 0 0 0 29.5 28.3 2000 0 0 0 29.1 28.0 10@09@08 0500 0 0 0 23.7 16.0 0600 0 0 0 23.7 16.0 0700 0 0 0 23.3 16.4 0800 0 0 0 23.3 16.8 0900 1 0 1 23.3 18.3 1000 1 1 2 23.3 20.6 1100 0 2 2 23.7 21.3 1200 2 0 2 24.4 21.7 1300 10 1 11 24.8 24.0 1400 15 0 15 25.6 24.8 1500 15 3 18 26.4 25.2 1600 6 1 7 26.8 24.0 1700 1 1 2 26.8 23.3 1800 0 0 0 26.8 23.7 1900 0 0 0 26.4 20.2 2000 0 0 0 26.0 19.0 10@10@08 0500 0 0 0 23.7 16.8 0600 0 0 0 23.3 16.4 0700 0 0 0 23.3 16.4 0800 0 0 0 23.3 17.1 0900 0 0 0 22.9 19.4 1000 1 0 1 23.3 22.5 1100 0 1 1 23.7 25.2 1200 5 0 5 24.4 26.8 1300 19 2 21 25.2 27.2 1400 16 2 18 26.0 28.3 1500 23 2 25 26.8 28.7 1600 13 0 13 27.2 28.3 29 Tabl 2.2Contd… 1700 5 1 6 27.2 27.2 1800 0 0 0 27.2 25.2 1900 0 0 0 27.2 22.1 2000 0 0 0 26.8 21.3 10@11@08 0500 0 0 0 24.0 18.3 0600 0 0 0 24.0 18.3 0700 0 0 0 24.0 18.3 0800 0 0 0 23.7 19.0 0900 0 0 0 23.7 22.5 1000 1 0 1 24.0 24.8 1100 2 0 2 24.4 26.8 1200 13 0 13 25.2 26.4 1300 19 2 21 26.0 28.3 1400 23 1 24 26.4 27.6 1500 25 2 27 26.8 28.0 1600 14 0 14 27.2 27.2 1700 2 0 2 27.2 25.6 1800 0 0 0 27.2 24.8 1900 0 0 0 26.8 23.3 2000 0 0 0 26.4 23.3
*Numb rsinboldd not th p akactivityduringth day
Discussion
Thisstudyisth firsttodocum ntdailyforagingactivityofL. bicolor inNorth
Am rica.Pr viousstudi sonth di lb haviorw r conduct dinPu rtoRicoand
Brazil.ForagingactivityofL. bicolor waspsinPu rtoRicowasb tw n0830and1530 hours(Castn randFowl r1987)notablyshort rthanth dailyforagingtim sr port d h r insumm r.Maximumactivityoccurswithsoilt mp ratur b tw n28@30°Candan airt mp ratur of30@35°C.Ev nthoughwasppopulationatn ctarsourc ss mtob mal bias d,g n rallyaround1300–1400hrs,mal :f mal ratiosonflow rsar n arly
30 on toon .Mal sandf mal sforag onflow rstoobtainn ctar(carbohydrat s)to sustainactivity.F mal shav amor ph m ralpr s nc du toactiv lys archingfor hostsforoviposition.Castn randFowl r(1987)alsor port dmal bias dpopulations onflow rs.Mal sw r n v robs rv dtoatt mpttomat whil f mal sforag dwhich isconsist ntwithobs rvationsbyM nk (1992)thatmatingoccursonth grass.Wasps w r obs rv dr stinginth shad dint riorofflow ringplantsandjustaf ww r obs rv dtosp ndth ov rnightp riodonth plant.R stinginth shad mayindicat a th rmor gulationb haviortocoolth thoracict mp ratur b caus suchb haviorwas notic dfrom1200hrsto1400hrswh nth t mp ratur wasatitsp akduringth day.
Waspl ftth plantsb tw n1800–1900hinsumm rand1700@1800hduringfall,butit isnotknownwh r th s waspssp ndth nightp riod.B caus airt mp ratur sdidnot s mtotrigg rthis,waspsmayus dusk(1845h)asatrigg rtol av th n ctarsourc s.
B caus th r isnopassiv monitoringsyst mforth wasp,dir ctobs rvations ar th onlym ansofmonitoring.Thisinpart xplainswhydatafroman ntir month w r pool dandnotpr s nt dass parat obs rvations.InJun 2007,for xampl ,th r wasa2w kp riodwh nnowaspsw r found.Thishasb ncall dth ‘summ r slump’(H.Frank,p rsonalcommunication)andd scrib sagapb tw nth m rg nc s oftwog n rations.St.Andr wsGolfCours r port dlyhadwaspsin2004(H ld2005) andsom w r not din2006.How v r,allsubs qu ntsurv ysfail dtoshowth wasps.
Th onlyr port dchang duringthistim isamor strictins cticid r gim whichis
vid ntinl ssmol crick tactivityonth cours .Th impactofturfgrassins cticid s onLarra bicolor hasnotb nstudi d.
31
CHAPTER III
ORNAMENTAL PLANTS AS NECTAR SOURCES FOR LARRA BICOLOR
Abstract
Parasiticwaspsg n rallyb n fitfromacc sstocarbohydrat s,primarilyinn ctar fromflow ringplants.Larra bicolor f donn ctarofc rtainwildflow rsandw ds includingSpermacoce verticillata.Th obj ctiv ofthisstudywasto valuat c rtain ornam ntalplantsasn ctarsourc sforadultL. bicolorandto valuat th pot ntial b n fitsofflow rf dingonlong vityandparasitism. PlantssimilartoSpermacocein structur andn ctarsourc w r us dtofindifanyofth mattract dLarra bicolor.A commongard n xp rim ntincluding13taxaofornam ntalplantsandS. verticillatain fourr plicat swasus dtod t rmin th pr f r nc ofL. bicolor forc rtainflow ring plantsund rfi ldconditions.Pentasattract dasmanywaspsasSpermacoce.Awhit @ flow r dvari tyofp ntaswasvisit dmor oft nbyL. bicolor thanoth rp ntas vari ti s.Th s r sultsw r furth rinv stigat dinlaboratorylong vityt sts.Wasp survivalonP ntaswascomparabl orb tt rthansurvivalofwaspsprovision dwith
Spermacoce.Th s r sultssugg stthatp ntasmayb plant dinlandscap stosustain thiswaspandp rhaps nhanc biologicalcontrol.
32
Introduction
Flow rf dingiscommoninparasitichym nopt rans(J rvis tal.1993)dir ctly
consumingfloraland xtrafloraln ctarorpoll n,toal ss r xt nttrichom s, pid rmis
orindir ctlyhon yd w.Pr s nc ofn ctaroroth rcarbohydrat sourc scanattractand
r tainnatural n mi s,(EvansandSwallow1993,JacobandEvans1998)th r by
incr asingparasitism.Incorporatingflow ringplantsadjac nttoturfgrasshas
implicationsforp stmanag m nt.Tiphia vernalis(Hym nop t ra:Tiphiida ),
introduc dforbiologicalcontrolofJapan s b tl (Popillia japonica)grubs,w r found
topr f rflow rsofPeonia spp.ingard nswithoth rbloomingplants(Pott rand
Rog rs2004).Inlandscap sorongolfcours s,d cr as dp standincr as dpr dator populationsw r obtain dwh ncons rvationstripsandwildflow rmix sw r
incorporat d(Braman tal.2002,FrankandShr wsbury2004).Similarly,parasitismof
Japan s b tl sgrubsbyTiphia vernalis wasgr at rwithin1mofapatchofflow ring
Peonia spp.(Pott randRog rs2004).
Similarpatt rnsar visibl in Larra bicolor,an ctoparasit of xotic
Scapteriscus mol crick ts,whichnotablyvisitsflow ringplantssuchasSpermacoce
verticillata (Rubiac a ).Th pr s nc ofn ctarsourc sar r port dlyimportantto
stablishm ntofL. bicolor(L ppla tal.2007).Arav loandFrank(2005) valuat d
foursp ci sofflow ringwildflow rs,Conoclinium coelestinum,Elephantopus elatus,
Passiflora coccinea,andSolidago fistulosaforvisitationsbyL. bicolor andsugarcont nt
33 ofth irn ctar.Allw r visit dl ssthanS. verticillata andhadalow rsucros @fructos ratio.
Flow ringornam ntalplantsp ci scommonlygrownfora sth ticsorfor attractingbutt rfli sar alr adypr s ntinurbanlandscap s.Id ntifyingflow ring plantsthatwouldalsoprovid an cologicals rvic mayfacilitat stablishm ntand succ ssofL. bicolorinlawnsandgolfcours turfgrass.Sinc S. verticillataisaw din coastalar as(Ev r st tal.1996)(Figur 3.2)andov rwint rsinth landscap incoastal
Mississippi(p rsonalobs rvations),th obj ctiv ofthisstudywastoinv stigat alt rnativ ornam ntalplantswithflow rsofsimilarshap andsiz asSpermacoce, particularlyoth rsp ci sinth Rubiac a .
InsouthMississippi,L. bicolor wasfoundf dingonPentas lanceolata
(Rubiac a )inth landscap (p rsonalobs rvationsFigur 3.1).Thissugg ststhat p ntasandp rhapsoth rflow ringplantsinth Madd rfamilycouldsubstitut asa n ctarsourc forL. bicolor.Vari ti sofP. lanceolatavaryinflow rcolorandplant h ight.P ntasar commoninsouth rnlandscap sforth irh attol ranc and attractiv n sstobutt rfli s(Brun r tal.2002).B caus p ntasar similarinflow r shap andinth sam familyasSpermacoce,oth rm mb rsofth Madd rfamilymay alsob attractiv toL. bicolor.
34
Figur 3.1
Larra bicolor F dingonPentas lanceolata
D spit th pr viousr s archonL. bicolor,th influ nc ofn ctarsourc son waspsurvivalorparasitismar notknown.L ppla tal.(2007)sp culat sthat“Th wasps(L. bicolor)willforag formol crick tsmor ff ctiv lyifS.verticillataplants ar availabl asasourc ofn ctar”.Thishypoth sisandth pr f r nc ofL. bicolor waspsforornam ntalplantsinth fi ldw r t st dtod t rmin th influ nc ofn ctar sourc sonwasplong vityinth laboratory.
35
Figur 3.2
Spermacoce verticillataasaW donth Roadsid sofFlorida
36 Materials and Methods
Source of Plants and Wasps
Plantsforthisstudyw r grownatth SouthMississippiBranchExp rim nt
StationinPoplarvill ,MSorpurchas dfromlocalr tail rs.Th plantsw r nottr at d withins cticid sbutw r f rtiliz dusing10@50@10(SchultzCo,Bridg ton,MO) r gularlytok pth mblooming.Waspsforus inth s xp rim ntsw r fi ld coll ct dwithin12hofth startofth xp rim nt.Waspsw r coll ct dusinga batt ry@pow r dvacuummodifi dforins ctcoll ction(#2820A,BioquipProducts,
RanchoDomingu z,CA)orcoll ct dusingana rialn t.Waspsw r stor dinacool r withic packsfortransporttoth lab.
Evaluation of Flowering Plants in a Common Garden Experiment.
InMay2007andin2008,agard nofpot ntialn ctarsourc swas stablish don th groundsofth CoastalR s archandExt nsionC nt r,Biloxi,MS.Eachplant sp ci sorcultivar(Tabl 3.1)was stablish dinfourr plicat dplots,0.5×0.5m.This
ntir gard nwascov r dwithabout5cmofgroundhardwoodmulchtor duc w d populationsanddripirrigationlin s stablish dthroughout.
B ginninginlat May achy ar,plantsw r insp ct dforL. bicolor. Dataw r
coll ct dwh nmor than2@3waspsw r inth gard n.Numb rsofwaspsonflow ring plantsw r r cord db tw n1100–1200h.Fift nminut obs rvationsw r mad in
achr plicat .Waspslandingonth flow rsw r count d v nifth ydidnotactiv ly
f d.Plantssp ci svisit dandth numb rofforagingwaspswasr cord dand
summariz dov r6din2007and10din2008.Numb rsw r th nanalyz dusing
37 PoisonAnalysistocompar th r lativ numb rofvisitstoc rtainornam ntalflow rs andd t rmin th pr f r nc ofLarra bicolor.Plotswh r plantsw r notflow ring w r xclud db for th analysis.
Tabl 3.1
HorticulturalPlantsEvaluat dasN ctarSourc sforLarra bicolor
Achillea millefolium Pentas lanceolata F nn l ‘Butt rflyPink’ Agastache urticifolium Pentas lanceolata Verbena ‘Azt c’ ‘Butt rflyWhit ’ Anethum graveolens Pentas lanceolata Verbena bonariensis ‘Butt rflyR d’ Cephalanthus Asclepias ‘SilkyGold’ Verbena ×hybrida occidentlis ‘Blu Lagoon’ Ixora coccinea Asclepias ‘SilkyR d’ Verbena ×hybrida ‘LadyScarl tt’ Lobularia maritime Bouvardia spp Verbena ×hybrida ‘QuartzWat rfall’
.
38
Figur 3.3
Th Gard nExp rim ntwithPlantsinBloom
Evaluation of Wasp Longevity in the Laboratory.
Laboratory xp rim ntsw r conduct don11–27August,2007tod t rmin th long vityofadultL. bicolorprovision dwithflow rs.Th s xp rim ntsw r conduct din22.7lit r,plasticcylindricalt star nas(St rilit ,Towns nd,MA,Figur
3.4).Th s ar nasw r 43.81cmtalltoaccommodat asmallpott dplant.Th bottom of achar awasfill dwithdrysandtoad pthof8cm.An mptyplasticpotwasplac d inth c nt rsothatplantscouldb asilyadd d.Asingl fi ld@coll ct dwaspwas introduc dto achar na.Inar naswh r flow ringplantsw r abs ntasugardi torno
39 di twasprovid d.Th sugardi twasa0.1%sugarsolutionwhichwasdabb dona
cottonballandplac donaplasticplatt r.Thiswaschang d v ryoth rdaytomak
sur thatsugarwat rwasavailabl .Waspsurvivalwas valuat ddaily.Th m an
numb rofdayswaspssurviv din achtr atm ntwasanalyz dusinganalysisofvarianc
(Statistix,AnalyticalSoftwar ,Tallahass ,FL)follow dbyTuk y’sm anss paration proc dur (P <0.05).
Figur 3.4
Ar naforth Long vityT sts
40
Results
Evaluation of Flowering Plants in a Common Garden Experiment.
Inbothy ars,L. bicolor waspsw r onlyobs rv donflow rsofPentasand
Spermacoce. Allvari ti sofp ntasr gardl ssofflow rcolorw r visit dbyL.
bicolor. Amongth s ,significantlymor waspsvisit dwhit @flow r dp ntasand
Sp rmacoc (Figur 3.5,F=12.09,P <0.0001).Abilityofr cruitingwaspswassimilar
forbothWhit P ntasandSpermacoce.
Figur 3.5
Fi ldT stR sultsofOrnam ntalPlantPr f r nc forLarra bicolor
41
Laboratory Longevity Assays.
Trial I: Waspsh ldincaptivitywithnofoodsurviv d,onav rag ,1.75dwhich
wasth sam asthos provision dwithflow ringSp rmacoc .Waspsprovision dwith
flow ringWhit P ntasliv d5dlong rthanthos provision dwithSp rmacoc (F=
10.12,P =0.012).Wasplong vityrang dfrom0–8daysinthis xp rim nt.
Figur 3.6
LabTrialI@Choic T stwithNoFoodControl,SpermacoceandPentas
Trial II: Th r w r nosignificantdiff r nc sb tw ntr atm ntsinthis
xp rim ntindicatingthatallthr foodsourc sar comparabl (F=1.1,P =0.39).
42 Wasplong vityrang dfrom2–6daysinthis xp rim nt.Th av rag long vityof mal sandf mal sw r 4.2and4.3d,r sp ctiv ly.
Figur 3.7
LabTrialIIwithSpermacoce,PentasandSugarWat rControl
Discussion
Flow rmorphology,color,orsugarcont ntofth n ctarmay xplainth similar r spons ofwaspstop ntasandSp rmacoc .Sp rmacoc isth primaryn ctarsourc forL. bicolor waspsinFlorida.Thissp ci sislist dasaw d(Ev r st tal.1996)and num rousvolunt rsofS. verticillata inar asadjac nttoth gard nplothav r quir d
43 ch micalcontrol.Fortunat ly,fi ldchoic t stssugg stthatanornam ntalsp ci s,
P ntas,mayb asuitabl r plac m nt.Th glossall ngthofmal andf mal L. bicolor ar notdiff r ntandroughly qualtoth flow rd pthofS.verticillata (Ar valoand
Frank2005).Inth fi ld xp rim nt,th r wasnodiff r nc innumb rsofwasps r cord donp ntaswithwhit flow rsandSp rmacoc .Flow rmorphology(i. .,siz ) lik lyhaslimit dimportanc sinc individualp ntasflow rsar larg rthanflow rsof
Sp rmacoc .Coloralon do snot xplainpr f r nc aswaspshav b nobs rv d f dingonplantswitharang offlow rcolors(Ar valoandFrank2005).
R sultsoflaboratorytrialsprovid additional vid nc thatPentasiscomparabl orp rhapsb tt rthanSpermacocer lativ towasplong vity.N ctarisan xc ll nt sourc ofcarbohydrat s(sugars)forins cts.Similarfi ldpr f r nc r s archwith
Pentasandbutt rfli s(Brun r tal.2002)alsosugg ststhatPentashasn ctarvolum or compositionwhichmak sitattractiv ton ctarf dingins cts.Waspflow rsand butt rflyflow rst ndtob sucros rich(Bak randBak r1983).Spermacocehas sucros @dominat dn ctar(H.Frank,unpublish ddata).D p@tub dflow rst ndtohav sucros richn ctar(Bak randBak r1983)andmay xplainth similarityforPentasand
Spermacoceinr cruitingwasps v nthoughSpermacocehassmallflow rscompar dto
Pentas.
Ev nthoughplantsinth sam familycouldshowsimilariti sinn ctarcont nt, thatn dnotb th cas inlarg famili s(Bak randBak r1983)andthis xplainswhy oth rplantsinth madd rfamilydidnotr cruitth wasp.Thisisindicationthatfloral charact risticslik floralvolatil sorn ctarcompositionwouldb th k ytowasp
44 pr f r nc .Sinc colorvariantsofth sam plantt ndtohav similarn ctarcont nt
(Bak randBak r1983)th abilitytor cruitwaspsbyth oth rcolorsofPentascouldb
xplain dasafunctionofth n ctarcont nt.Th diff r nc inwaspattractiondu to
colorn dnotb du todiff r nc inn ctarcont nt.
45
CHAPTER IV
FIRST REPORT OF LARRA BICOLOR (HYMENOPTERA: SPHECIDAE) IN
ALABAMA
Abstract
Larra bicolorisaparasiticwaspintroduc dintoFloridaforbiologicalcontrolof p stmol crick ts.Waspshuntformol crick tsinth pastur sandmanag dturfgrass.
Onc th waspfindsamol crick t,th ychas ittoth surfac andstingsit.Th sting
t mporarilyparalyz sth mol crick t nablingth wasptod positan gg.Th wasp
gghatch sandlarvaf dsasan xt rnalparasit .Th larvaconsum sth mol crick t
th nspinsacocooninth soil.Th r ar 2@3g n rationsofL. bicolor achy ar.Wasps hav spr adthroughoutFloridaandintoth north rnGulfstat ssuchasMississippi wh r L. bicolor w r discov r din2004.Th pr s ntstudywasconduct dtod t rmin ifL. bicolor waspr s ntongolfcours sinMobil andBaldwinCounti s(south rnmost counti s)inAlabama.Infall2007, ightgolfcours sinth s counti sw r id ntifi d andsurv y dforL. bicolor.Waspsw r obs rv dorcoll ct don4ofth 8golfcours s surv y d.Th discov ryofL. bicolor insouth rnandcoastalAlabamaprovid san w r cordforthissp ci sbutmor importantlyitprovid sn winformationforturfgrass manag rs.On@goingr s archproj ctsar inv stigatingth rol offlow ringplantsin
46 parasitismandsurvivalofL. bicolor.Turfgrassmanag rsandhom own rsinAlabama cannowutiliz th findingsofthisandfutur r s archonbiologicalcontrolofmol crick ts.
Introduction
Mol crick ts(Orthopt ra:Gryllotalpida :Scapteriscusspp.)ar s riousp stsof warms asonturfgrassandpastur sinth south rnUnit dStat s.Sinc th ir ntryinto
Unit dStat salongth G orgiacoastlin inth arly1900’s,Scapteriscus mol crick ts hav spr adsouthintoFloridaandw stthroughLouisiana.InMississippiandth north rnGulfr gion,twosp ci s,S. vicinus andS. borellei,ar pr s ntwh r asth s twoandS. abbreviatus ar pr s ntinFlorida.Damag fromScapteriscusmol crick ts ismosts v r inth south rnmostar asofGulfstat s(FrankandWalk r2006).Ongolf cours s,ins cticid sar us d xt nsiv lytopr v ntdamag frommol crick ts.For
xampl ,controlmaycostupto$675p rha.Combin d ff ctsofthr natural n mi s,
Ormia depleta(Dipt ra:Tachinida ),Steinernema scapterisci( ntomopathog nic n matod ),andLarra bicolorF.(Hym nopt ra:Sph cida )ar attribut dwitha99% r ductioninmol crick tpopulationsinFlorida(FrankandWalk r2006).
47 Figur 4.1.
a. Larra bicolor(15–20mmlong) isShownH r F dingonFlow rsof Spermacoce verticillata b.StingingaMol Crick t.
Larra bicolor (Figur .4.1)isan ctoparasitoidofmol crick tsandhas2to3 g n rationsp rs ason(FrankandSourakov2002).Th biologyand cologyofL. bicolor w r r c ntlyr vi w d(Frank tal.1995)andwillnotb discuss dind tail.
Larra bicolor waspsar blackwithwingsthatar browntoblu andasolidr dabdom n
(Figur .4.1)whichdistinguish sitfromLarra analis F.whichisnativ toMississippi andoth rGulfstat s,andhasablackabdom nwithr djustatth tip(M nk 1992).
B tw n1981–1983,r s arch rscoll ct dL. bicolor waspsfromPu rtoRicoand r l as dth matfiv sit sinFlorida.Followingth s r l as s,waspsonlyb cam
stablish datth south rnmostr l as sit n arFt.Laud rdal (Frank tal.1995).Inan att mptto stablishpopulationsinnorth rnFlorida,thr sp ci sofwasps,Larra bicolor,L. praedatrix,andL. godmaniw r coll ct dfromBoliviaandr l as dalong withparasitiz dhostsatthr sit sn arGain svill in1988@89(Frank tal.1995).
48 Again,th s r l as sapp ar dunsucc ssfuluntil1993wh nL. bicolor w r obs rv d attackingmol crick tsandf dingonflow rsofSpermacoce verticillata inGain svill .
In2004, s v ralsp cim nsofL. bicolor w r coll ct dfromgolfcours sin coastalMississippi.Larra bicolor curr ntlyiswid spr adinFlorida(Frank tal.1995,
FrankandWalk r2006)andr port dfromsouth rncounti sinMississippiandG orgia
(H ld2005).Th mostnorth rncoll ctionofL. bicolor inMississippiis(30.69°N)
Sauci r,MS(D.W.H ld,unpublish ddata)andmost stablish dpopulationsofL. bicolor inth U.S.occurat≤31°Nlatitud (FrankandWalk r2006).Larra bicolor was int ntionallyr l as dinsouth rnG orgiain2000byW.Hudsonforbiologicalcontrolof mol crick ts(W.Hudson,p rsonalcommunication).Withnor cordsofr l as in
Mississippi,th discov ryofL. bicolor sugg stsnaturalrang xpansionofthiswasp alongwithitshost(H ld2005).Rang xpansionfromFloridatoMississippiwould indicat thatL. bicolor mayalsob pr s ntinAlabama.Forthisr ason,w sampl dgolf cours sinth south rnandcoastalar asofAlabamatod t rmin ifandwh r Larra bicolor waspr s nt.Iffound,turfmanag rsinsouthAlabamawouldb n fitfrom knowingL. bicolor ispr s ntandcouldutiliz on@goingr s archto nhanc local populationsandparasitism.
Materials and Methods
Forthissurv y,south rnAlabama(<31°Nlatitud )wasdivid dintothr r gions;th m troMobil ar a,th ast rnshor ofMobil Bay,andcoastalAlabama includingGulfShor s.B tw n26S ptand19Oct.2007,golfcours sin achr gion w r sampl dforL. bicolor.On achcours ,th gr nsstafforsup rint nd nts
49 sugg st datl astthr fairwaysinf st dwithmol crick tstosampl .Surv ysw r
conduct db tw n0800–1400hours.Allobs rv rswalk d,sid bysid ,inalin
spanningth widthofth fairway.Ev ryobs rv rwalk d achfairwayfromt to
gr n.Particularatt ntionwasgiv ntoar aswithfr shmol crick tdamag .IfL.
bicolor waspsw r obs rv datt mptsw r mad tocoll ctth m.Wh ncaptur d,wasps
w r coll ct dwithn tsth ntransf rr dtocoll ctionbottl sinacool rfortransportto
th laboratory.Ifnotcaptur d,th waspwasnot dasobs rv dbutnotcoll ct dandth
surv yr sum dtoth ndofth fairway.
Results and Discussion
Larra bicolor waspsw r foundonatl aston golfcours in achofth thr g ographicalr gionsinsouthAlabama(Tabl 4.1).Thisr cordisth firstr portofthis b n ficialwaspinAlabama.Itisprobabl thatth populationsofL. bicolor stablish d insouthAlabamahav r sult dfromnaturalrang xpansionofthiswaspwithinits host’srang (H ld2005).
AdultL. bicolor wasrar lyfoundinallar assurv y donaparticulargolfcours .
Waspsw r mor commoninar asorongolfcours swh r fr shmol crick tdamag wasmor appar nt.Som golfcours sw r mor aggr ssiv withth irmol crick t manag m ntprogramsandthos cours shadf w rorsmall rar asthatw r damag d.
B caus L. bicolorwaspsar difficulttosampl ,non@d t ctionofwaspsonc rtainsit s duringthissurv yshouldnotb quat dwithanabs nc ofwaspsonthatcours or g ographicalar a.Inasurv yofgolfcours sincoastalMississippiin2004,H ld(2005)
50 r port dc rtainsit swh r L. bicolorw r pr s ntorabs nt.Sinc thatsurv y,L. bicolor hasb ncoll ct dfromon sit wh r itwasund t ct din2004,and populationsar nowabs ntfromon sit wh r th yw r onc pr s nt(H ldand
Abraham,unpublish ddata).T mporalors asonalabundanc inwasppopulationsis lik lyd t rmin dbyhostabundanc ,pr s nc ofn ctarsourc s(L ppla tal.2007),or surfac @appli dins cticid s.
Tabl 4.1
Numb rsofLarra bicolor R port dfromGolfcours sinSouthAlabama.
Golf course Observed Collected
Metro Mobile (Mobile Co.) SpringHillColl g 9 3 CountryClubofMobil 0 0 Azal aCity 3 1 Eastern side of Mobile Bay (Baldwin Co.) Timb rCr k 0 0 QuailCr k 12 4 Coastal (Gulf Shores and Orange Beach) KivaDun s 0 0 P ninsula 8 0 GolfClubatth Wharf 0 0
51
Figur 4.2 a.AF mal L. bicolor Pr paringtoEnt rth MoundinS archforaMol Crick t.b. Mol Crick tMoundwithHol thatwasCommonlyObs rv donSit swh r L. bicolor Waspsw r Activ .
Onthr ofth foursit swh r L. bicolor waspsw r found,mol crick t moundswithhol sinth topofth mw r alsoobs rv d(Figur .4.2a,b).Larra bicolor f mal smov alongth grasssurfac s archingformounds.Onc amoundis d t ct d,th soilisant nnat dandth waspmaychoos to nt r.To nt ramound,th f mal mustburrowthroughth soilatth surfac .Itth nproc dsintoth subt rran an tunn lsins archofahost.Asar sult,mol crick tscommonly xitth moundtoavoid attack.Hol ssimilartoth s ar commonlyobs rv dons v ralsit sinsouth
Mississippiwh r L. bicolor ar pr s nt.Th s hol s,th r for ,mayr sultfrom ith r th wasp nt ringorth mol crick t scapingth moundandmayb apassiv indicationthatL. bicolor isactiv inthatar a.
52 Th discov ryofL. bicolor insouth rnandcoastalAlabamaprovid san w
r cordforthissp ci sbutmor importantlyitprovid sn winformationforturf
manag rs.On@goingr s archproj ctsinFlorida(H.Frank,p rsonalcommunication)
andAlabamaar inv stigatingth rol offlow ringplantsinparasitismandsurvivalof
L. bicolor.Turfmanag rsinAlabamacannowutiliz th findingsofthisandfutur r s archonbiologicalcontrolofmol crick ts.
53
CHAPTER V
CONCLUSIONS
D t ctionofLarra bicolorinMississippiin2004indicat dthatth wasp
xpand ditsrang fromFloridawithoutanyhumanint rf r nc .Oursubs qu nt d t ctionofL. bicolor inAlabamasupportsthisconc ptofnaturalrang xpansion.In som Alabamasit s,w obs rv dgr at rnumb rsofwaspsthanwhatar s ninon locationinMississippi,whichsugg stpopulationsmayhav first stablish dinAlabama.
On ofth r asonsattribut dtoth succ ssofth wasp stablishm ntinFlorida hasb nth availabilityofitsn ctarsourc Spermacoce verticillata whichisaw din south rnturf.Fi ldandlabstudi sw conduct dindicat thatPentascouldsustainth waspforlong rtim sthanSpermocace.Thisprovid sanoth rr asontoinclud Pentas inourlandscap atl astwh r mol crick tsar aprobl m.
S asonalactivityinth north rngulfr gionis noughfor2@3g n rations,and willb actingonasingl g n rationofmol crick tsthat m rg inay ar.Th di l activitythoughwasmor int r sting,wh nitshow dtwodistinctpatt rns,wh nth y sporadicallyapp ar donth n ctarsourc sduringsumm randwh nth yhadap ak activityp riodinth fallwh nth t mp ratur sr ach dth maximuminth dayb for th iractivityagainc as daroundsuns t.Th s obs rvationsindicat thatth activityof th waspsisd p nd ntonacombinationofsunris ,suns ttimingsandt mp ratur s.
54 Withth informationthisstudyprovid s,w ar b tt r quipp dtoutiliz th s rvic sofLarra bicolorasabiocontrolag ntasw knowthatthatth yar availabl in south rnpartsofMississippiandAlabama.W knowthatPentascansubstitut
Spermacoceasan ctarsourc andwithknowl dg aboutitsdailyands asonalactivity,
w couldtim ourins cticid applicationslat rinth day,and v nr duc partofitto
low rth costsofcontrolandalsotoaugm ntwasppopulations.
55
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