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Billbug (Sphenophorus spp.) Composition, Abundance, Seasonal Activity, Development Time, Cultivar Preference, and Respon...

Permanent Link: http://ufdc.ufl.edu/UFE0022248/00001

Material Information

Title: Billbug (Sphenophorus spp.) Composition, Abundance, Seasonal Activity, Development Time, Cultivar Preference, and Response to Endophytic Ryegrass in Florida
Physical Description: 1 online resource (71 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: billbug, cultivar, endophyte, golf, sphenophorus, turfgrass
Entomology and Nematology -- Dissertations, Academic -- UF
Genre: Entomology and Nematology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Billbugs (Sphenophorus spp.) are common pests whose damage is often misdiagnosed on turfgrass in the United States. Florida turfgrass managers experiencing billbug outbreaks have been struggling to obtain satisfactory control using conventional insecticide applications. Thus, I have sought to identify the billbug (Sphenophorus spp.) complex, seasonal activity, and development time in golf course bermudagrass, document billbug responses on bermudagrass and zoysiagrass cultivars, and determine the effect of endophytic perennial ryegrass on billbug development. Adult billbug populations were monitored weekly from January 2006 to December 2007 with four linear pitfall traps on each of two bermudagrass golf courses in southern Florida and two in north-central Florida. Several studies of bermudagrass and zoysiagrass cultivar resistance were conducted in the greenhouse by caging adult Sphenophorus venatus vestitus onto plastic pots in 2006 and 2007. The total number of notches, eggs, and larvae were recorded to evaluate resistance. Overseeding of endophytic perennial ryegrass into bermudagrass was conducted in the greenhouse and in the field to evaluate the impact to S. venatus vestitus oviposition and larval development. Four endoyphytic perennial ryegrass cultivars were also tested for S. venatus vestitus feeding damage and oviposition potential. My results suggested that S. venatus vestitus is the most abundant billbug species on Florida golf courses containing bermudagrass, constituting > 80% of the specimens collected from the 24-hr samples, and the remainder consists of nine other Sphenophorus spp. As many as 686 individuals of S. venatus vestitus were collected within a 24-hr sampling period during peak adult activity in April 2006 on one golf course in southern Florida, but activity was steady on the other three courses ( < 150 adult billbugs per 24-hr sample). Sphenophorus venatus vestitus shows greater preference for ?Tifway? bermudagrass than other cultivars, and Zoysia matrella cultivars display higher resistance than cultivars of Zoysia japonica. Bermudagrass overseeded with endophytic perennial ryegrass significantly reduced S. venatus vestitus feeding damage and oviposition.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Buss, Eileen A.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0022248:00001

Permanent Link: http://ufdc.ufl.edu/UFE0022248/00001

Material Information

Title: Billbug (Sphenophorus spp.) Composition, Abundance, Seasonal Activity, Development Time, Cultivar Preference, and Response to Endophytic Ryegrass in Florida
Physical Description: 1 online resource (71 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: billbug, cultivar, endophyte, golf, sphenophorus, turfgrass
Entomology and Nematology -- Dissertations, Academic -- UF
Genre: Entomology and Nematology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Billbugs (Sphenophorus spp.) are common pests whose damage is often misdiagnosed on turfgrass in the United States. Florida turfgrass managers experiencing billbug outbreaks have been struggling to obtain satisfactory control using conventional insecticide applications. Thus, I have sought to identify the billbug (Sphenophorus spp.) complex, seasonal activity, and development time in golf course bermudagrass, document billbug responses on bermudagrass and zoysiagrass cultivars, and determine the effect of endophytic perennial ryegrass on billbug development. Adult billbug populations were monitored weekly from January 2006 to December 2007 with four linear pitfall traps on each of two bermudagrass golf courses in southern Florida and two in north-central Florida. Several studies of bermudagrass and zoysiagrass cultivar resistance were conducted in the greenhouse by caging adult Sphenophorus venatus vestitus onto plastic pots in 2006 and 2007. The total number of notches, eggs, and larvae were recorded to evaluate resistance. Overseeding of endophytic perennial ryegrass into bermudagrass was conducted in the greenhouse and in the field to evaluate the impact to S. venatus vestitus oviposition and larval development. Four endoyphytic perennial ryegrass cultivars were also tested for S. venatus vestitus feeding damage and oviposition potential. My results suggested that S. venatus vestitus is the most abundant billbug species on Florida golf courses containing bermudagrass, constituting > 80% of the specimens collected from the 24-hr samples, and the remainder consists of nine other Sphenophorus spp. As many as 686 individuals of S. venatus vestitus were collected within a 24-hr sampling period during peak adult activity in April 2006 on one golf course in southern Florida, but activity was steady on the other three courses ( < 150 adult billbugs per 24-hr sample). Sphenophorus venatus vestitus shows greater preference for ?Tifway? bermudagrass than other cultivars, and Zoysia matrella cultivars display higher resistance than cultivars of Zoysia japonica. Bermudagrass overseeded with endophytic perennial ryegrass significantly reduced S. venatus vestitus feeding damage and oviposition.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Buss, Eileen A.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0022248:00001


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1 BILLBUG ( Sphenophorus spp.) COMPOSITION, ABUND ANCE, SEASONAL ACTIVITY, DEVELOPMENT TIME, CULTIVAR PREFER ENCE, AND RESPONSE TO ENDOPHYTIC RYEGRASS IN FLORIDA By TA-I HUANG A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2008

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2 2008 Ta-I Huang

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3 To my dear parents

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4 ACKNOWLEDGMENTS I sincerely thank Dr. Eileen A. Buss, m y s upervisory committee chair, for her support and guidance and for giving me an opportunity for my professional and personal growth. She has been a role model of professiona l honesty, work ethic, enthusiasm and dedication balanced with love and care in the personal life, which will inspire me throughout my further career. Working with Dr. Buss was enjoyable and rewarding. I wa nt to acknowledge my supervisory committee members (Dr. Marc Branham and Dr. Kevin Kenwor thy) for their contribution. I am grateful for the research sites, assistance and cooperation provided by Mark Kann (University of Florida Plant Science Unit in Citra, FL), Michael Rowe (Gainesville Golf a nd Country Club), Mark Dickson (WestEnd Country Club), Frank Sbarro (LaGorce Country Club), Sean Anderson (Card Sound Golf Club), Dr. Leah Brilman from Seed Re search of Oregon who provided me with seed product, and the Scotts Company provided me the ry egrass seeds. I want to thank all of my lab partners who helped me with my research: Cara Vazquez, Olga Kostromytska, Jessica Platt, Paul Ruppert, Jade Cash, Megan Gilbert, and Rachel Sheahanand. I also would like to thank Dr. Michael Thomas (Division of Plant Industries) and Lyle Buss (University of Florida) for their help. I am grateful for the funding provided by the Florida Turfgrass Association, Florida Golf Course Superintendents Associati on, and Golf Course Superintendents Association of America.

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5 TABLE OF CONTENTS Page ACKNOWLEDGMENTS...............................................................................................................4LIST OF FIGURES.........................................................................................................................8ABSTRACT.....................................................................................................................................9CHAPTER 1 INTRODUCTION..................................................................................................................11Warm Season Grasses in North America............................................................................... 11Common Arthropod Pests of Bermudagrass..........................................................................11Billbugs Sphenophorus spp. in the United States................................................................... 12Sphenophorus venatus vestitus in the Southeastern United States......................................... 13Billbug Integrated Pest Management...................................................................................... 142 BIOLOGY AND SEASONAL PH E NOLOGY OF BILLBUGS, Sphenophorus spp., IN FLORIDA...............................................................................................................................16Materials and Methods...........................................................................................................17Composition, Abundance, and Seasonal Activity of Sphenophorus spp. in Florida....... 17Sphenophorus venatus vestitus Adult Activity Patterns, Fecundity and Generation Time.............................................................................................................................18Adult daily activity patterns..................................................................................... 18Potential fecundity and egg development of S. venatus vestitus ..............................19Variation of adult S. venatus vestitus body size....................................................... 20Length of S. venatus vestitus Development in Bermudagrass and Zoysiagrass.......20Larval instar determination...................................................................................... 21Results.....................................................................................................................................21Composition, Abundance, and Seasonal Activity of Sphenophorus spp. in Florida....... 21Sphenophorus venatus vestitus Adult Activity Patterns, Fecundity and Generation Time.............................................................................................................................23Adult daily activity patterns..................................................................................... 23Potential fecundity and egg development of S. venatus vestitus ..............................23Variation of adult S. venatus vestitus body size....................................................... 24Length of S. venatus vestitus development in bermudagrass and zoysiagrass.........24Larval instar determination...................................................................................... 25Discussion...............................................................................................................................253 EVALUATION OF BEMUDAGRASS AND Z OYSIAGRASS RESISTANCE TO Sphenophorus venatus vestitus ...............................................................................................42Materials and Methods...........................................................................................................43Damage potential of adult S. venatus vestitus on Fur Brmudagrass Cltivars.................. 43

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6 Plant material............................................................................................................ 43Adult damage potential............................................................................................ 43Bermudagrass Cltivar Peference Test............................................................................. 44Plant material............................................................................................................ 44Preference evaluation............................................................................................... 44Zoysiagrass Cultivar Resistance to S. venatus vestitus ...................................................44Plant material............................................................................................................ 44Resistance evaluation...............................................................................................45Results and Discussion......................................................................................................... ..46Damage Potential of Adult S. venatus vestitus on Four Bermudagrass Cultivars........... 46Bermudagrass Cultivar Preference Test.......................................................................... 46Zoysiagrass Cultivar Resistance to S. venatus vestitus ...................................................474 EFFECT OF ENDOPHYTE LEVEL IN PERENNIAL RYEGRASS ON THE SURVI VAL AND DEVELOPMENT OF Sphenophorus venatus vestitus ............................56Materials and Methods...........................................................................................................57Sphenophorus venatus vestitus Survival, Development, and Damage on Four Endoyphytic Perennial Ryegrass Cultivars.................................................................. 57Impact of Overseeding Two Bermudagra ss Cultivars with Endophytic Perennial Ryegrass on S. venatus vestitus Survival, Development, and Damage.......................57Overseeded Bermudagrass Field Trial............................................................................ 58Results and Discussion......................................................................................................... ..59Sphenophorus venatus vestitus Survival, Development, and Damage on Four Endoyphytic Perennial Ryegrass Cultivars.................................................................. 59Impact of Overseeding Two Bermudagra ss Cultivars with Endophytic Perennial Ryegrass on S. venatus vestitus Survival, Development, and Damage.......................59Overseeded Bermudagrass Field Trial............................................................................ 60APPENDIX A EVALUATION OF BEMUDAGRASS RESISTANCE TO Sphenophorus inaequalis ........63Materials and Methods...........................................................................................................63Damage Potential of Adult S. inaequalis on Four Bermudagrass Cultivars................... 63Plant material............................................................................................................ 63Adult damage potential............................................................................................ 63Results and Discussion......................................................................................................... ..64Damage Potential of Adult S. inaequalis on Four Bermudagrass Cultivars................... 64LIST OF REFERENCES...............................................................................................................66BIOGRAPHICAL SKETCH.........................................................................................................71

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7 LIST OF TABLES Table Page 2-1 Total number of Sphenophorus spp. collected from four linea r pitfall traps at each of four Florida golf courses during 24-hour sam ple periods from January 2006 to December 2007.................................................................................................................. 30 2-2 Mean (SEM) number of eggs per female S. venatus vestitus collected from three golf courses in Florida in 2007.......................................................................................... 31 2-3 Mean (SEM) number of eggs per female S. inaequalis collected at W est End Country Club in 2007.........................................................................................................32 2-4 Mean ( SEM) body lengths of male and female S. venatus vestitus at three golf courses in 2007. ............................................................................................................... ..33 3-1 Damage between male and female S. venatus vestitus on four berm udagrass cultivars in the greenhouse...............................................................................................................50 3-2 Mean (SEM) number of notches and surviving larvae per pot on four bermudagrass cultivars in the greenhou se preference test. .......................................................................51 3-3 Ratings (SEM) of seventeen zoysia grass cu ltivars afte r 4 weeks of adult S. venatus vestitus infestation.............................................................................................................. 52 3-4 Mean number (SEM) of S. venatus vestitus eggs and larvae, a nd characteristics of adult notches on 17 zoysiagrass cultivars, after 1 month of adult confinem ent................ 53 3-5 Mean number (SEM) of S. venatus vestitus eggs and larvae, and adult notches on 18 zoysiagrass cultivars, after one m onth of adult confinement........................................ 54 4-1 The effect of overseeding with endophy tic pe rennial ryegrass on two bermudagrass cultivars on S. venatus vestitus damage potential and oviposition....................................62 A-1 Damage between male and female S. inaequalis on f our bermudagrass cultivars in the greenhouse...................................................................................................................65

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8 LIST OF FIGURES Figure Page 2-1 Total number of S. venatus vestitus a dults collected weekly from four linear pitfall traps at Gainesville Country Club....................................................................................344 2-2 Total number of adult S. venatus vestitus obtained weekly from f our linear pitfall traps at LaGorce Country Club........................................................................................ 355 2-3 Total number of adult S. venatus vestitus trapped weekly at C ard Sound Country Club..................................................................................................................................366 2-4 Total number of S. inaequalis adu lts collected each week from four linear pitfall traps at West End Country Club...................................................................................... 377 2-5 Mean number of S. venatus vestitus adults th at were activ e on the soil surface of bermudagrass pots during a 24hour period of observation.............................................389 2-6 Number of life stage for S. venatus vestitus reared on pots of Tifway bermudagrass in the greenhouse. ..............................................................................................................39 2-7 Number of life stage for S. venatus vestitus reared on pots of Em pire zoysiagrass in the greenhouse...................................................................................................................40 3-1 Sphenophorus venatus vestitus preference test for berm udagrass cultivars.................... 555

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9 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science BILLBUG ( Sphenophorus spp.) COMPOSITION, ABUND ANCE, SEASONAL ACTIVITY, DEVELOPMENT TIME, CULTIVAR PREFER ENCE, AND RESPONSE TO ENDOPHYTIC RYEGRASS IN FLORIDA By Ta-I Huang May 2008 Chair: Eileen A. Buss Major: Entomology and Nematology Billbugs ( Sphenophorus spp.) are common pests whose da mage is often misdiagnosed on turfgrass in the United States. Florida turfgrass managers e xperiencing billbug outbreaks have been struggling to obtain satisfact ory control using conve ntional insecticide applications. Thus, I have sought to identify the billbug ( Sphenophorus spp.) complex, seasonal activity, and development time in golf course bermudagra ss, document billbug responses on bermudagrass and zoysiagrass cultivars, and determine the e ffect of endophytic perenni al ryegrass on billbug development. Adult billbug populations were monitored weekly from January 2006 to December 2007 with four linear pitfall traps on each of two bermudagrass golf courses in southern Florida and two in north-central Florida. Several studies of bermudagrass and zoysiagrass cultivar resistance were conducted in the greenhouse by caging adult Sphenophorus venatus vestitus onto plastic pots in 2006 and 2007. The total numbe r of notches, eggs, and larvae were recorded to evaluate resistance. Overseeding of endophy tic perennial ryegrass into bermudagrass was conducted in the greenhouse and in the field to evaluate the impact to S. venatus vestitus oviposition and larval

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10 development. Four endoyphytic perennial ry egrass cultivars were also tested for S. venatus vestitus feeding damage and oviposition potential. My results suggested that S. venatus vestitus is the most abundant billbug species on Florida golf courses containing bermudagrass, constituting >80% of the specimens collected from the 24-hr samples, and the remainder consists of nine other Sphenophorus spp. As many as 686 individuals of S. venatus vestitus were collected within a 24-hr sampling period during peak adult activity in April 2006 on one golf course in southern Florida, but activity was steady on the other three courses (<150 adult billbugs per 24-hr sample). Sphenophorus venatus vestitus shows greater preference for Tifway berm udagrass than other cultivars, and Zoysia matrella cultivars display higher resistance than cultivars of Zoysia japonica Bermudagrass overseeded with endophytic perennial ryegrass significantly reduced S. venatus vestitus feeding damage and oviposition.

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11 CHAPTER 1 INTRODUCTION Warm Season Grasses in North America War m-season turfgrasses are widely grown in th e transition zone and southern part of the United States. These typical warm s eason turfgrasses include bahiagrass ( Paspalum notatum Fluegge), bermudagrass (Cynodon dactylon (L.) Pers), centipedegrass (Eremochloa ophiuroides (Munro) Hack.), seashore paspalum ( Paspalum vaginatum Sw.), St. Augustinegrass ( Stenotaphrum secundatum (Walt.) Kuntze.), and zoysiagrass ( Zoysia spp.). Bermudagrasses are among the most widely used warm-season grasses on golf courses, athletic fields, and in highprofile residential and commercial landscapes where a fine-textured, dense ground cover is desired (Trenholm 2003). Tif way, Tifdwarf, and Tifeagle are the most popular bermudagrass cultivars used on golf course roughs, fairways, and greens in Florida (Russ Meyer, personal communication 2006). With over 1,500 golf courses, Florida turfgrass managers and golfers demand high quality turf. However, turfgra ss quality can be impaired by insects that feed on grass leaves, stems, sap, or roots (Buss 2003). Common Arthropod Pests of Bermudagrass Common arthropod pests that feed on be rm udagrass include mole crickets ( Scapteriscus spp.), several caterpillar species, bermudagrass mites [ Eriophyes cynodoniensis (Sayed)], white grubs (Coleoptera: Scar abaeidae), and billbugs ( Sphenophorus spp.). Mole crickets damage turfgrass by tunneling in the soil, uprooting plants and feeding on turfgras s roots (Buss et al. 2006). Several species of caterpillars [e.g., Spodoptera spp., Mocis spp.] skeletonize and completely consume grass leaf blades, causing an uneven and off-colored appearance to the turfgrass (Buss and Turner 2004). Feeding by the bermudagrass mite results in shortened stems and stolons, yellow and curly blades, and tuft s of grass plants (Short and Buss 2005). White

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12 grubs (especially masked chafers, Cyclocephala spp.) feed on fine turf grass roots, which makes bermudagrass turn slightly yellow, but they also are food for birds that di g in the turf to find them (Buss and Turner 2004). Billbug larvae are stem borers and root feeders, and adults can damage stems, but their damage is often misdiagnosed as drought stress or disease pressure. Infested grass turns yellow and results in pa tchy spots during a heavy infestation (Brandenburg and Villani 1995). Billbugs Sphenophorus spp. in the United States Although over 60 native billbug sp ecies exist, rou ghly nine species have been reported to damage turfgrass in North America: the bluegrass billbug, S. parvulus Gyllenhal; the lesser billbug, S. minimus Hart; S. venatus Say (no common name); the uneven billbug, S. inaequalis Say; the hunting billbug, S. venatus vestitus Say; the Phoenician billbug, S. phoeniciensis Chittenden; the Denver billbug, S. cicatristriatus Fabraeus; S. coesifrons Gyllenhal; and S. apicalis LeConte (Vaurie 1951, Tashiro 1987, Morrill and Suber 1976, Johnson-Cicalese et al. 1990). Sphenophorus parvulus is the most common pest of c ool-season turf, and attacks Kentucky bluegrass, perennial ryegrass, fine fescue and tall fescue from the Paci fic Northwest across to New England (Tashiro 1987). It ha s also been reported in several of the Gulf States where warmseason turfgrasses predominate (Richmond 2000). Sphenophorus minimus occurs in the northeastern U.S., feeding on cool-season turf grasses. The life cycle and host range of S. minimus is very similar to S. parvulus (Johnson-Cicalese et al. 1990, Shetlar 1991). The distribution of S. inaequalis is limited to the eastern United St ates (Veurie 1951). It attacks both warm and cool season turfgrasses including be rmudagrass, Kentucky bl uegrass, perennial ryegrass, and tall fescue. The life cycle of S. inaequalis is similar to S. parvulus and its population could be as abundant as S. parvulus in New Jersey (Johnson-Cicalese and Funk

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13 1990). The most common billbug in warm season turfgrass is S. venatus vestitus It occurs north to New Jersey and west to Texas, and is mo st abundant in and damaging to bermudagrass and zoysiagrass. Adult females chew holes or notches in the grass stems where they deposit their eggs (Brandenburg and Villani 1995). The distribution of S. phoeniciensis is limited to southern California and Arizona. Its life cycle and hosts are similar to S. venatus vestitus (Tashiro 1999). Sphenophorus cicatristriatus occurs in the Rocky Mountain re gion from northern New Mexico to Nebraska and prefers feeding on cool seas on turfgrasses such as Kentucky bluegrass and perennial ryegrass (Vaurie 1951, Tashiro 1987, Shetlar 1995). Sphenophorus cicatristriatus overwinters as a late instar larva, and th e adult population peaks between June and midSeptember (Vaurie 1951, Shetlar 1995). The life cycle of both S. coesifrons and S. apicalis are poorly understood. The distribution of S. coesifrons extends throughout the southern United States while S. apicalis occurs from New Jersey to th e Gulf States (Vaurie 1951). Sphenophorus venatus vestitus in th e Southeastern United States The species considered most abundant and dama ging in the southeastern United States is S. venatus vestitus. Adults (6-11 mm long) cause notches while feeding on stem tissue. Legless larvae are usually white to yellowish with brown head capsules. First instar larvae feed inside stems and then drop out into the soil. Mature la rvae feed on roots in soil and pupate in a chamber about 2 to 5 cm deep (Brandenburg and Villani 1995). Sphenophorus venatus vestitus occurs from New Jersey to Florida and along the Gulf states west to Texas. It is also present in Mexico and some of the Caribbean islands, and was tran sported with sod to the Middle East, Southeast Asia and Hawaii (Vaurie 1951, Tashir o 1999, Niemczyk and Shetlar 2000). The most commonly recorded hosts in Florida are bermudagrass and zoysiagrass. Other minor hosts include bahiagrass, centipedegrass, St. Augustinegrass, orchardgrass ( Dactylus glomeratus L.), crabgrass ( Digitaria spp.), signal grass ( Brachiaria decumbens Stapf),

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14 barnyardgrass ( Echinochloa crusqalli Beauv.), corn ( Zea mays L.), sugarcane ( Saccharum officinarum L.), and leather leaf fern ( Polypodium scouleri Hook and Grev.) (Satterthwait 1919, Kelsheimer 1956, Kamm 1969, Oliver 1984). Satterthwait (1931) listed some additional hosts of S. venatus vestitus: yellow nutsedge ( Cyperus esculentus L.), wheat ( Triticum aestivum L.), timothy (Phleum pratense L.), and a bulrush ( Scirpus validus Vahl.) (Woodruff 2005). Billbug Integrated Pest Management Billbugs are considered one of the m ost misd iagnosed pests of turfgr ass (Potter 1998). Turf managers usually misdiagnose billbug damage sy mptoms as drought stress, delayed green up in the spring, disease such as dollar spot, or ot her insect feeding damage (Buss 2006). Billbug infestations may be managed using resistant tu rfgrasses, overseeding with endophyte-enhanced turfgrasses, biopesticides and pr eventive or curative in secticides. Asay et al. (1983) reported a genetic resistance of range grasses to S. parvulus Later, several studie s demonstrated endophyteenhanced resistance in perennial ryegrass to S. parvulus and other Sphenophorus spp. (Ahmad et al. 1986, Jonhson-Cicalese and White 1990, and Richmond 2000). Little is known about the biological control of billbugs, but their na tural enemies include parasitoids, nematodes, and fungi. Satte rthwait (1919) reported a parasitic wasp, Vipio belfragei Cresson, reared from billbug larvae, and a mymarid egg parasitoid, Anaphoidea calendrae Gahan, was documented to parasitize several Sphenophorus spp. (Satterthwait 1931). Several entomopathogenic nematodes (e.g., Steinernema carpocapsae S. feltae and Heterorhabditis bacteriophora ) suppress both white grubs and bill bugs (Niemczyk and Shetlar 2000). Muscardine fungi, Beauveria spp., were found to attack severa l billbug species in New Jersey (Johnson-Cicalese 1988), but thei r effectiveness is unknown. A comprehensive knowledge of the billbug life cy cle and seasonal phenology is critical for insecticidal control. A preven tive application in spring for billbug control is recommended by

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15 most extension reports and turfgrass management websites. The goal is to kill adults emerging in the spring before they lay eggs, or kill first in stars as they emerge and begin feeding. Early season applications to manage S. parvulus have been effective in western New York lawns (Tashiro and Personius 1970). However, regions with multiple, overlapping generations and a prolonged adult activity period may have difficulty controlling infest ations only with insecticides. An integrated pest management (IPM) program needs to be developed for billbugs on warmseason turfgrasses. Given the limited information about genera l billbug biology and management in warm season turfgrasses in Florida, I sought to desc ribe the species composition, abundance, seasonal activity, developmental time of Sphenophorus spp. on golf courses utilizing bermudagrass (Chapter 2), document billbug preference on bermuda grass and zoysiagrass cultivars (Chapter 3), and billbug response to endophytic perennial ryegrass (Chapter 4).

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16 CHAPTER 2 BIOLOGY AND SEASONAL PH E NOLOGY OF BILLBUGS, Sphenophorus spp., IN FLORIDA Over 60 native billbug species exist in the United States (Niemczyk and Shetlar 2000), and at least nine of those species attack turfgrass (V aurie 1951, Morrill and Suber 1976, Johnson-Cicalese et al. 1 990). The bluegrass billbug, Sphenophorus parvulus Gyllenhal, typically infests cool season turfgrasses in th e northeastern United St ates (Tashiro 1987), including Kentucky bluegrass ( Poa pratensis L.), perennial ryegrass (Lolium perenne L.), tall fescue ( Festuca arundinacea Schreb.), and 14 other non-turf grasses (Johnson-Cicalese and Frank 1990). The lesser billbug, S. minimus Hart, also infests cool season grasses in the northeastern United States (J ohnson-Cicalese et al. 1990, Shetlar 1991). The uneven billbug, S. inequalis Say, occurs in the eastern United States on Ke ntucky bluegrass, pere nnial ryegrass, tall fescue and bermudagrass ( Cynodon dactylon (L.) Pers.) (Satterthw ait 1931, Johnson-Cicalese and Frank 1990). Little is known about S. apicalis LeConte, but it occurs in the Gulf States and New Jersey (Vaurie 1951). Sphenophorus coesifrons (no common name) is distributed throughout the southern United States (V aurie 1951), and at tacks bahiagrass ( Paspalum notatum Flgg.) (Morrill and Suber 1976). The Denver billbug, S. cicatristriatus Fahraeus, occurs in the Rocky Mountain region from northern New Mexico to Nebraska and prefers feeding on cool season turfgrasses such as Ke ntucky bluegrass and perennial ryegrass (Vaurie 1951, Tashiro 1987, Shetlar 1995). The Phoenician billbug, S. phoeniciensis (Chittenden), infests warm season grasses in southern California and Arizona, and the orchardgrass billbug, S. venatus confluens Chittenden, is found in Oregon. Its primary host is orchardgrass (Dactylus glomeratus L.) (Kamm 1969). The hunting billbug, S. venatus vestitus Chittenden, infests primarily bermudagrass and zoysiagrass ( Zoysia spp.) from New Jersey to Florida and along the Gulf States (Tashiro 1987).

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17 Billbugs are both stem boring and root feed ing turfgrass insects (Potter 1998). Adult females chew holes or notches into grass stem s that are utilized for egg deposition (Brandenburg and Villani 1995). Eggs (about 1.5 mm long) are cl ear to creamy white, and hatch in 3-10 days depending on the temperature (Potter 1998). Young larvae (about 1.3 mm long) tunnel in and hollow out stems, and leave sawdust-like frass inside the stems (Potte r 1998). Mature larvae (about 6-10 mm long) severely damage grass by f eeding on crowns, stolons, and roots. Pupae (about 1.3-1.5 mm long) occur in a soil chambe r 2.5-5.1 cm deep (Shetlar 1995). Adults (8-11 mm in length) have elongate snouts, elbowed an tennae, and hard elytra (Brandenburg and Villani 1995), and different species may be identified by unique pronotal patterns (Vaurie 1951, Johnson-Cicalese et al. 1990). Although 25 species of billbugs occur in Florida (Peck and Thomas 1998), only S. venatus vestitus is credited as being the most abundant an d damaging species. The seasonal activity of S. venatus vestitus is poorly known in Florida, and although it has only one generation per year in northern states, it has been speculated that several overlapping generations occur along the Gulf States (Potter 1998). My goals here include 1) Determining which S phenophorus spp. are most abundant in northern and southe rn Florida 2) Determining the development time and seasonal phenology of S. venatus vestitus and S. inaequalis Materials and Methods Composition, Abundance, and Seasonal Activity of Sphenophorus spp. in Florid a The composition, abundance, and seasonal activity of Sphenophorus spp. on four golf courses in Florida were determined weekly by taking 24-hour samples with large linear pitfall traps from January 2006 to December 2007. The gol f courses included Gainesville Country Club and West End Country Club in Gainesville, LaGorce Country Club in Miami Beach, and Card

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18 Sound Country Club in Key Largo, Fl orida. Four linear pitfall tr aps (similar to Lawrence 1982) were placed in bermudagrass ( Cynodon dactylon (L.) Pers.) roughs on each golf course in January 2006. Each trap had four 3-m long arms of PVC pipe (7.6 cm diameter), with a 2.5 cm slit cut lengthwise in each arm. The far pipe end was capped, a nd the inner pipe end was placed in a hole on the side of a 19 liter bucket. A removable plastic tube extended the arm over a removable 4 liter bucket. Rocks were placed be neath the 19 liter bucket, and both buckets had holes drilled in the bottom to allow water drainage To sample, debris was removed from traps in the morning, sand was added to the bottom of the 4 liter bucket, and any billbugs caught during the following 24-hours were collected and pres erved in 70% EtOH. Adult billbug species, abundance and gender were determined for each sampling date. Weather conditions for each sampling period were recorded. The soil activity of Sphenophorus spp. was examined at the same golf courses monthly from 24 January 2006 to 15 December 2006. Four cores (10.2 cm diameter, 15 cm deep) were collected near damaged areas on bermudagrass roughs. Cores were placed in plastic bags, transported to the laboratory, and dissected under at least 10X magni fication. Any billbug life stages present were collected, counted, and pr eserved in 70% EtOH. This sampling method was discontinued in 2007 because few specimens were recovered. Sphenophorus venatus vestitus Adult Activity P atterns, Fecundity and Generation Time Adult daily activity patterns The diurna l activity patterns of adult S. venatus vestitus were observed every 4 hours each day on 30 April 2007 (15 pots), 11 November 2007 (10 pots), 15 January 2008 (10 pots), and 7 February 2008 (10 pots) (7 observation intervals). Two male and two female S. venatus vestitus adults were placed onto each Tifway bermuda grass pot (11.4 cm diameter). Adults were allowed to acclimate for 2 days before observatio ns began. The adults in each pot were observed

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19 for 20 seconds, and the location of adults (e.g., gra ss blade, soil surface, not visible) and their behavior (e.g., grooming, walking, feeding, mating, or inactive) were recorded (but not timed). Red light was used for night observations to minimize disturbance. Weather conditions were recorded at each observation period. Data were analyzed using two-way ANOVAs (SAS Institute 2000) to detect the effect of time and date on billbug activity. Means were compared using Tukeys HSD test ( P < 0.05). Potential fecundity and egg development of S. venatus ves titus Adult female S. venatus vestitus collected from the linear pi tfall traps at the four golf courses from 22 January to 27 December 2007 were dissected with a 10X dissecting microscope. Twenty or more females were dissected at leas t monthly from each site The number of mature eggs (1.5 to 1.7 mm long) in th eir abdomens was recorded. The time of ovarial development at each golf course, and whether or not females were in reproductive diapause were determined by examination of the ovaries (Young 2002). Data were analyzed using two-way ANOVAs (SAS Institute 2000) to detect the eff ect of month and site on female S. venatus vestitus potential fecundity. Means were compared using Tukeys HSD test ( P < 0.05). Twenty female S. venatus vestitus were collected from pitfall traps at Gainesville Country Club on 9 August 2006 and placed in a container (15 15 cm) with moistened filter paper. Females readily laid eggs in the absence of pl ant tissue. Twenty-five newly laid eggs (<24 hrs old) were collected from the container and pl aced in a Petri dish (8.8 cm diameter) with moistened filter paper in a rearing room under 24-26C, 25% re lative humidity, light intensity of ~4.05 lum/m2, and a photoperiod of 10:14 (L:D) hrs. The percentage of eggs hatching each day was determined until all eggs had hatched or died.

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20 Variation of adult S. venatus vestitus body si ze Twenty male and 20 female adult S. venatus vestitus that were collected from the pitfall traps at Gainesville Golf and Country Club, LaGorce Country Club, and Card Sound Country Club each in March, July, and November 2006 were examined for differences in body size. The body length from the base of the beak to the tip of the anus was measured with a 10X dissecting microscope and ocular micrometer. Data were analyzed using a two-way ANOVA (SAS Institute 2000) to detect the effect of time and site on S. venatus vestitus body length. Means were compared using Tukeys HSD test ( P < 0.05). Length of S. venatus ve stitus Development in Bermudagrass and Zoysiagrass Thirty plastic pots (11.4 cm diameter) were pl anted with cores of Tifway bermudagrass on 30 March 2007, and 15 pots each of Empire and P ristine zoysiagrass were similarly potted on 4 May 2007, and allowed to establish in the gr eenhouse for one month. All cores were obtained from the University of Floridas Plant Science Res earch Unit in Citra, FL. Pots were fertilized with 113.5 g of Miracle-Gro all purpose fertilizer (20-20-20) per week and irrigated daily. Sixty pairs of adult S. venatus vestitus were collected on Tifway bermudagrass on 27 April 2007 and on Empire zoysiagrass on 19 June 2007 at the U.F. Plant Science Research Unit in Citra, FL. Two pairs of S. venatus vestitus were randomly chosen and ca ged onto each pot within 24 hours of collection. Each pot was encircled up to 15 cm above the turf height wi th a clear plastic tube to prevent adult escape. All adults were rem oved from pots five days after introduction. Six different pots were randomly selected and destruct ively sampled to find all billbug life stages at 3, 6, 9, 11, and 12 weeks post-adult removal in the bermudagrass trial and every 2 weeks in the zoysiagrass trial (2, 4, 6, 8, 10 weeks post-adu lt removal). Larvae were preserved in KAAD for 48 hours and stored in 70% EtOH. Air temperature, relative humidity, and soil temperature were recorded in the greenhouse throughout the experiment.

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21 Larval instar determination To estim ate the number of instars for S. venatus vestitus, the head capsule widths of 135 larvae reared on bermudagrass were measured under a 10X dissecting microscope equipped with an ocular micrometer. First instar head capsule widths were measured on larvae that hatched from eggs that females laid in a Petri dish (8.8 cm diameter) with moistened filter paper. Results Composition, Abundance, and Seasonal Activity of Sphenophorus spp. in Florid a From January 2006 to December 2007, a total of 18,580 adult billbugs of ten different Sphenophorus spp. were collected from the f our golf courses (Table 2-1). Sphenophorus venatus vestitus was the most abundant specie s collected, constituting 80.9% of all specimens (Table 21). Gainesville Golf and Country Club and West End Country Club had the greatest species diversity, each with eight sp ecies collected during the 24hour sampling periods. LaGorce Country Club had the lowest species di versity, with three species collected. Sphenophorus venatus vestitus was the primary species (99.9% of all specimens) collected from the two southern Florida golf courses. The sex ratio of all the samples combined was 1.1:1 (males to females) for S. venatus vestitus, 1.9:1 for S. inaequalis and 1.2:1 for S. cariosus Olivier. Billbug gender was determined by examining the ventral si de of the abdomen; males had a full, rounded abdomen, and female abdomens were depressed. In addition to the specimens collected during the 24-hour sampling period, six additional Sphenophorus spp. were found when traps were cleaned, including S. callosus Olivier, S. coesifrons Gyllenhal, S. minimus Hart, S. necydaloides Fabricius, S. pontederiae Chittenden, and S. dietrich Satterthwait (a new Alachua County record). Sphenophorus inaequalis is a new Monroe County record and S. costipennis Horn is a new Florida state record (Peck and Thomas 1998).

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22 Adult S. venatus vestitus were collected from pitfall traps nearly every week of the year. The timing of peak trap catches of S. venatus vestitus varied on each golf course. At Gainesville Golf and Country Club, it was most abundant on 7 March 2006 (68 adul ts per 4 traps), at LaGorce Country Club on 10 April 2006 (685 adults ), and at Card Sound Country Club on 4 July 2007 (163 adults) (Figures 2-1 to 23). The months of greatest overall S. venatus vestitus adult activity were March and April. Specimens were not collected on several da tes because traps were flooded or cooperators at the tw o southern golf courses were unable to check them. Given the variability of adult S. venatus vestitus activity over time and locati on, the number of generations per year could not be determined through pitfall trapping. The second most abundant species collected was S. inaequalis constituting 17.8% of all specimens (Table 2-1). It was collected at three of the four golf courses, but was most abundant at West End Country Club. Peak abundance of S. inaequalis at West End Country Club was on 17 July 2006 with 145 adults co llected during the 24 hour sampli ng period. The mean number of S. inaequalis adults collected by month was highest in July 2006 (108.5 12.7) and lowest in February 2006 (13.2 7.2) (Figure 2-4). No eggs or young larvae were found from turf in the cupcutter samples collected in 2006. Nearly mature larvae were only found in the soil. With the results from all four golf courses combined, four larvae were collected in March, three in April, one in May, four in July, and one in October 2006. One pupa was recovered in Marc h. One adult was collected in April, one in May, two in June, one in July, one in September, and two in Oct ober. Species identification of immature Sphenophorus spp. was not possible.

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23 Sphenophorus venatus vestitus Adult Activity P atterns, F ecundity and Generation Time Adult daily activity patterns Adults were seen feeding, m ating, walking, or inactive at all observation times, but were most active on the soil surface between midnight and 4:00 am ( F = 52.79; df = 5, 314; P < 0.0001). The mean number of adults observed on each date was also significantly different ( F = 26.82; df = 3, 314; P < 0.0001). Mating and grooming were only observed between midnight and 4:00 am (20% and 5% of adults, respectively). During observations air temperature ranged from 18.6 29.5 C, soil temperature ranged from 63 72C, and relative humidity ranged from 40 86%. The mean number of adults observed on the surface between midnight and 4:00 am on 1 May 2007 ( F = 13.18; df = 6, 104; P < 0.0001), 11 November 2007 ( F = 8.95; df = 6, 69; P < 0.0001), 15 January 2008 ( F = 5.21; df = 6, 69; P = 0.0005), and 7 February 2008 ( F = 20.05; df = 6, 69; P < 0.0001) were all significan tly higher than any other time period (Figure 2-5). The interaction between each observation time and date was not significantly different. Potential fecundity and egg development of S. venatus ves titus The potential fecundity of S. venatus vestitus varied by location and time of year. Mature eggs were found from female ovaries almost ever y month of the year on each golf course except for June at LaGorce Country Club and November at Card Sound Country Club. Number of mature eggs per female were 0 to 14 (4.9 0.1) at the Gainesville Golf and Country Club, zero to 14 (4.9 0.1), 0 to 9 (3.1 0.2) at LaGorce Count ry Club, and 0 to 9 eggs (3.4 0.16) at Card Sound Country Club (Table 2-2). Fr om all data combined, female S. venatus vestitus collected from Gainesville Golf and Country Club containe d significantly more mature eggs than both golf courses in southern Florida ( F = 13.5; df = 2, 33; P < 0.0001). Fecundity was greatest in March at LaGorce Country Club (F = 5.03; df = 10, 233; P < 0.0001) and in October at Gainesville Golf and Country Club ( F = 2.88; df = 11, 654; P = 0.0011). From the Petri dish rearing, 44 % (11

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24 eggs) of the eggs hatched 7 days after oviposition and 40% (10 e ggs) after 8 days. The remaining four eggs died within 6 days of oviposition. At West End Country Club, the number of mature eggs found in female S. inaequalis ovaries ranged from 0 to 5 (1.9 0.05) (Table 2-3). Sphenophorus inaequalis females contained significantly more mature eggs during the months of March, July and September than other months ( F = 6.52; df = 11, 669; P < 0.0001). Variation of adult S. venatus vestitus body si ze The average body length of female S. venatus vestitus (8.3 0.03 mm; range: 7.1 to 9.6 mm) was significantly greater than for males (7.5 0.03 mm; range: 6.6 to 8.4 mm) (F = 320.69; df = 1, 359; P < 0.0001), regardless of time or location. Body lengths of males and females collected from Gainesville Golf and Country Club were significantly greater than for males and females collected from LaGorce Country Club and Card Sound Country Club (male: F = 8.05; df = 2, 179; P = 0.0005) (female: F = 15.51; df = 2, 179; P < 0.0001) (Table 2-4). Body lengths of all S. venatus vestitus collected in March, July, and Novemb er combined, did not significantly differ among golf courses. Length of S. venatus ve stitus development in bermudagrass and zoysiagrass Bermudagrass. From 24 May to 26 July 2007, a total of two eggs, 45 larvae, seven pupae, and 17 adults were found during the five eval uations. At the first time evaluation, 24 May 2007, only two eggs and ten larvae were found (two larvae were in the soil and the rest were in the stems). Thirty-one larvae and only one pupa were found on 14 June 2007. Three larvae, three pupae and two adults were found on 5 July 2007. On e larva, three pupae, a nd seven adults were found on 19 July 2007. Eight adults were found on 26 July 2007 (Figure 2-6). Zoysiagrass. From 7 July to 3 September 2007, a total of five eggs, 24 larvae, two pupae, and eight adults were found in th e five evaluations. Five eggs and seven larvae were found on 9

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25 July 2007. Five larvae were found on 23 July 200 7. Seven larvae and one pupa were found on 6 August 2007. Five larvae, one pupa, and one a dult were found on 20 August 2007. Seven adults were found on 3 September 2007 (Figure 2-7). Larval instar determination The head capsule width of S. venatus vestitus larvae ranged f rom 0.4 to 2.4 mm. The head capsule width of first instars was about 0.5 0.02 mm (n = 20), and the head capsule width of late instar larvae was about 2.3 0.02 mm (n = 15). The number of instars could not be determined, but 4-5 instars may exist, based on the frequency distribut ion (Figure 2-8). Discussion Despite the potential for a larg e species complex, S. venatus vestitus was the most abundant species present on bermudagrass in th e four golf courses monitored. It was also expected to be the most damaging species, given its reputation in warm season turfgrass (Tashiro 1987), but distinct damage was only visible on LaGorce Country Club, which had the highest number of adult S. venatus vestitus collected throughout the study (Table 2-1). The composition of the billbug species comple x varied between northern and southern Florida and by golf course. Speci es composition may vary in the state due to different horticultural zones, soil types, or manage ment practices, but Ti fway bermudagrass was commonly grown at all four locations and be rmudagrass is widespread on golf courses throughout the state. Sphenophorus venatus vestitus has a broad geographic range in North America, spanning both cool and warm seas on grasses (Johnson-Cicales e et al. 1990). Young (2002) also reported similar numbers of S. venatus vestitus in Arkansas. However, JohnsonCicalese et al. (1990) reported th at the four billbug species ( S. inaequalis, S. minimus S. parvulus, and S. venatus vestitus) occurred nearly equal numbers in New Jersey. In my study, S.

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26 minimus and S. parvulus were only found in Gainesville a nd their populations were smaller compared to S. venatus vestitus. The life cycle of S. venatus vestitus and body size appears to al so vary with location, but may be more greatly influenced by temp erature or host plant. In general, S. venatus vestitus overwinters as adults in norther n states, adult population increase s in spring, then declines in summer, another smaller population increases in the fall and decrease s in winter (JohnsonCicalese et al. 1990, Young 2002). It has one generation per year in northern states (Tashiro 1987), and may have a partial second generation in New Jersey (Johnson-Cicalese et al. 1990). It is also suspected to have two generations in Georgia (S. K. Braman, personal communication). The development from oviposition to adult emergence in my greenhouse study at ca. 25.8 C took ca. 10-11 weeks on bermudagrass and ca. 27 C took ca. 8-10 weeks. In addition, female S. venatus vestitus develop mature eggs every month of th e year and can live at least one month. Therefore, it is possible that S. venatus vestitus has at least 2 to 3 ove rlapping generations each year in Florida. Another similar species, S. venatus confluens, overwinters as adults, begins to feed in March, and adults die in July as larv ae emerge and feed throughout the summer (Kamm 1969). However, the seasonal activity from my pitfall trapping data in Florida showed different adult activity by different locations. Sphenophorus venatus vestitus adults were most abundant in March and April. The population then decrease d during the summer followed by a smaller peak again during late fall. Difference in climate may affect in the activity patterns between southern and northern Florida. The pitfall trap data are difficult to interpret because of the variability in infestation levels among the golf courses sampled. Adult activity pattern of Sphenophorus spp. within a 24-hour period is fairly unknown. An observation study on the southern corn billbug, S. callosus indicated that adults were most active

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27 during daylight hours (DuRant 1985). Activity was mo re associated with air temperature than with soil temperature, and was inhib ited when air temperature was below 20oC (DuRant 1985). However, adult observations in this study showed an opposite activity peri od from the previous study. Adult S. venatus vestitus activity was greater at midnigh t than any other timing, which suggests that S. venatus vestitus adults may be nocturnal. Although some behavior was observed during the day, most adults emerged on the soil surface to feed and mate at night. However, billbug activity may last until early morning since many of Sphenophorus spp. are observed on sidewalk in early morning (OBri en, 2006 personal communication). Johnson-Cicalese et al. (1990) repor ted a long oviposition period of S. venatus vestitus from spring to early September in New Jersey. In Arkansas, ovarian development of S. venatus vestitus was quickly mature in early April, a nd remained highly mature until October (Young 2002). In Oregon, S. venatus confluens deposit a large number of eggs in the stems and on the leaf sheaths in mid-June (Kamm 1969). In this study, the number of eggs deposited in grass could not be determined from golf courses, so the precise time of ovarial maturation was determined by dissecting females from weekly tr ap collections. In general, mature eggs of S. venatus vestitus were found every month in 2007, but were higher in early spring and late fall than in summer. Mature eggs were still presented in December which may suggest a second generation occurred in Florida, and S. venatus vestitus overwinters in both adult and immature stages. The reason why the fecundity of S. venatus vestitus in Gainesville was significantly higher than in southern Florida is unknown. One possi ble explanation may be the difference in body length. Since a lot of species of Coleoptera have been proved than female size is a principal constraint on insect potential fecundity (Hone k 1993), the significantly longer body length of S.

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28 venatus vestitus in Gainesville (Table 2-4) resulting in higher fecundity than in south Florida is reasonable. The other possibility may be popul ation density. Reigada and Godoy (2005) reported that fecundity and body size are generally density -dependent characters influenced by insect populations. Female S. venatus vestitus in Gainesville have to produ ce more eggs because of its low population and greater species di versity; In contrast, female S. venatus vestitus in south Florida may not need high fecundity since the population is large and there is less species competition (Table 2-1). In greenhouse rearing studies, adult billbugs were caged into plastic pots by encircling with clear plastic tubes 15 cm above the turf height. This met hod proved excellent for preventing adults from escaping. Since flying activity wa s only observed in a container twice for S. venatus vestitus and several times for S. inaequalis I speculate that crawli ng is the major method of population dispersal. Young (2002) reported that flight occurs sporadically by only a few S. venatus vestitus during spring, and only less than 10 % of the dissected adults developed flight muscle. Although adult S. venatus confluens have fully developed wings and make short flights 2-3 feet high in May and Octobe r in late afternoon (Kamm 1969) he believes walking is the usual mode of spread in the field. Hansen (1987) divided the instars of S. parvulus into three classes by larval head capsule width: first instars < 0.6 mm, middle instars between 0.6 and 1.15 mm, and late instars < 1.15 mm. Four S. venatus confluens larvae reared on orchardgrass in the greenhouse had 5 instars (Kamm 1969). Although there is no direct evidence shows the number of instars of S. venatus vestitus from my larval head capsule data, at least 4 to 5 instars was speculated. The second most abundant billbug species was the uneven billbug, S. inaequalis It occurs throughout the eastern United States and west to Texas, and nort h to central Florida (Peck and

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29 Thomas 1998). The life cycle of S. inaequalis is barely known. JohnsonCicalese et al. (1990) found several S. inaequalis in the summer, but the population was more abundant in spring and autumn in New Jersey, and was nearly equal in abundance with three other billbug species. These result differ from our sampling data in Florida where there was a large adult S. inaequalis population peak from mid-June to late August 2006 at West End Country Club. It is not clear why the S. inaequalis population in 2007 was le ss than 2006, but there was still an obviously consistent population peak from June to Augus t in 2007 which indicated one generation per year of S. inaequalis The known hosts of S. inaequalis are bermudagrass, Kentucky bluegrass, perennial ryegrass and fescues (Satterthw ait 1931, Johnson-Cicalese and Funk 1990). JohnsonCicalese et al. (1990) suggeste d an equal pest status of S. inaequalis as the other three species in New Jersey. However, a large number of S. inaequalis was only collected at West End Country Club which suggest it may only cause damage in localized area of Florida. Adult S. venatus vestitus can be found every month throughout the year in Florida with an extended oviposition period by females. The multiple overlapping generations makes control more difficult compared to the univoltine blue grass billbug in northern states. However, among the golf courses studied in Florida, serious bi llbug damage was only observed at one (LaGorce Country Club in Miami Beach). This might indi cate that most well managed bermudagrass turf can tolerate certain levels of billbug attac k. However, the irregularly distribution of Sphenophorus spp. and overlapping life cycles at differe nt locations in Florida need further studies. This study provides a fundamental knowledge of billbug species complex and seasonal activity in Florida for pest management.

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30 Table 2-1. Total number of Sphenophorus spp. collected from four lin ear pitfall trap s at each of four Florida golf courses during 24-hour sample periods from January 2006 to December 2007. Golf Course Species Gainesville West EndLaGorceCard SoundTotal S. apicalis 62 44 4 1 111 S. cariosus 5 25 0 0 30 S. cubensis 1 0 3 1 5 S. deficient 0 2 0 0 2 S. inaequalis 13 3,331 0 1 3,345 S. minimus 38 4 0 0 42 S. necydaloides 1 0 0 0 1 S. parvulus 0 2 0 0 2 S. pontederiae 1 1 0 0 2 S. venatus vestitus 2,073 428 8,898 3,641 15,040 Total 2,194 3,837 8,905 3,644 18,580

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31 Table 2-2. Mean (SEM) number of eggs per female S. venatus vestitus collected from three golf courses in Florida in 2007. Gainesville Country Club LaGorce Country Club Card Sound Country Club Month No. females No. eggs No. femalesNo. eggsNo. females No. eggs January 11 5.6 0.8 15 3.3 0.9 15 4.2 0.8 February 22 3.7 0.6 16 3.6 0.6 13 3.0 0.7 March 35 5.7 0.4 18 5.1 0.5 18 4.7 0.6 April 89 5.5 0.3 23 3.3 0.5 20 2.4 0.5 May 62 4.5 0.4 20 1.8 0.4 31 3.7 0.3 June 62 4.4 0.4 --20 3.2 0.4 July 88 4.3 0.3 20 1.8 0.4 22 3.0 0.5 August 81 4.8 0.3 20 3.4 0.6 20 3.2 0.5 September 56 4.8 0.3 20 2.6 0.3 20 3.0 0.4 October 63 6.0 0.4 20 3.2 0.4 20 3.4 0.5 November 71 5.1 0.4 42 4.2 0.3 --December 16 3.6 0.5 20 4.1 0.4 20 4.8 0.4

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32 Table 2-3. Mean (SEM) number of eggs per female S. inaequalis collected at West End Country Club in 2007. Means within columns with different letters are statistically different at = 0.05 ( F = 6.52; df = 11, 669; P < 0.0001) Month No. females Mean no. eggs January 10 2.0 0.3 February 21 1.8 0.2 March 31 2.3 0.2 April 70 1.3 0.1 May 48 1.3 0.1 June 95 1.9 0.1 July 92 2.3 0.1 August 90 1.8 0.1 September 60 2.4 0.2 October 70 2.0 0.1 November 46 1.4 0.2 December 25 1.3 0.2

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33 Table 2-4. Mean ( SEM) body lengths of male and female S. venatus vestitus at three golf courses in 2007. Golf Course Male Body length max.min. (mm) Female Body length max.min. (mm) Mean # male body length (mm) Mean # female body length (mm) Gainesville 6.8 8.3 7.8 9.5 7.7 0.04a 8.5 0.1a LaGorce 6.6 8.4 7.3 9.6 7.5 0.10b 8.3 0.1b Card Sound 6.8 8.4 7.1 9.3 7.5 0.04b 8.1 0.1b Means within columns with different letters are statistically different at = 0.05 ( F = 8.05; df = 2, 179; P = 0.0005) Means within columns with different letters are statistically different at = 0.05 ( F = 15.51; df = 2, 179; P < 0.0001)

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34 0 10 20 30 40 50 60 7024-Jan 7-Feb 21-Feb 8-Mar 21-Mar 5-Apr 18-Apr 2-May 16-May 31-May 20-Jun 6-Jul 18-Jul 2-Aug 22-Aug 6-Sep 19-Sep 3-Oct 17-Oct 31-Oct 14-Nov 28-Nov 15-Dec 2-Jan. 16-Jan 30-Jan 16-Feb 27-Feb 14-Mar 27-Mar 11-Apr 25-Apr 8-May 23-May 5-Jun 20-Jun 4-Jul 17-Jul 31-Jul 14-Aug 30-Aug 12-Sep 25-Sep 10-Oct 24-Oct 7-Nov 21-Nov 5-Dec 19-Dec2006 2007# S. venatus vestitus collected Figure 2-1. Total number of S. venatus vestitus adults collected weekly fro m four linear pitfall traps at Gainesville Country Club.

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35 0 100 200 300 400 500 600 7007-Feb 21-Feb 7-Mar 21-Mar 4-Apr 24-Apr 3-May 24-May 13-Jun 29-Jun 12-Jul 26-Jul 8-Aug 22-Aug 19-Sep 17-Oct 7-Nov 20-Nov 6-Dec 20-Dec 9-Jan 28-Jan 13-Feb 12-Mar 26-Mar 10-Apr 10-amy 13-Jun 11-Jul 17-Aug 28-Aug 17-Sep 23-Oct 21-Nov 18-Dec2006 2007# S. venatus vestitus collecte d Figure 2-2. Total number of adult S. venatus vestitus obtained weekly from four linear p itfall traps at LaGorce Country Club.

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36 0 20 40 60 80 100 120 140 160 1807-Feb 21-Feb 8-Mar 21-Mar 5-Apr 18-Apr 9-May 23-May 13-Jun 26-Jul 8-Aug 23-Aug 19-Sep 24-Oct 15-Nov 28-Nov 12-Dec 26-Dec 3-Jan 25-Jan 31-Jan 19-Feb 12-Mar 5-Apr 19-Apr 21-May 5-Jun 26-Jun 11-Jul 17-Aug 17-Sep2006 2007# S. venatus vestitus collected Figure 2-3. Total number of adult S. venatus vestitus trapped weekly at Card Sound Country Club.

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37 0 20 40 60 80 100 120 140 16024-Jan 7-Feb 21-Feb 8-Mar 21-Mar 5-Apr 18-Apr 2-May 16-May 31-May 20-Jun 6-Jul 18-Jul 2-Aug 22-Aug 6-Sep 19-Sep 3-Oct 17-Oct 31-Oct 14-Nov 28-Nov 15-Dec 2-Jan. 16-Jan 30-Jan 16-Feb 27-Feb 14-Mar 27-Mar 11-Apr 25-Apr 8-May 23-May 5-Jun 20-Jun 4-Jul 17-Jul 31-Jul 14-Aug 30-Aug 12-Sep 25-Sep 10-Oct 24-Oct 7-Nov 21-Nov 5-Dec 19-Dec2006 2007# S. inaequalis collecte d Figure 2-4. Total number of S. inaequalis adults collected each week from four linear pitfall trap s at West End Country Club.

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38 0 0.5 1 1.5 2 2.5 3 3.5 4 4:00pm8:00pm12:00am4:00am8:00am12:00pm4:00pmMean # of S. venatus vestitus observed 04/30/07' 11/11/07' 01/15/08' 02/07/08' Figure 2-5. Mean number of S. venatus vestitus adults that were active on the soil surface of bermudagrass pots during a 24hour period of observation.

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39 0 5 10 15 20 25 30 35 5/24/20076/14/20077/5/20077/19/200707/26/2007Number of S. venatus vestitus collecte d # Eggs # Larvae # Pupae # Adults Figure 2-6. Number of life stage for S. venatus vestitus reared on pots of Tifway bermudagrass in the greenhouse.

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40 0 1 2 3 4 5 6 7 8 07/09/200707/23/200708/06/200708/20/200709/03/2007Number of S. venatus vestitus collected # Eggs # Larvae # Pupae # Adults Figure 2-7. Number of life stage for S. venatus vestitus reared on pots of Empire zoysiagrass in the greenhouse.

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41 0 2 4 6 8 10 12 14 16 18 0.00.51.01.52.02.53.0 Head capsule width (mm)Number of larva e Figure 2-8. Frequency di stribution of head capsule width (mm) of S. venatus vestitus larvae collected from greenhouse rearing and eggs.

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42 CHAPTER 3 EVALUATION OF BEMUDAGRASS AND Z OYSIAGRASS RESISTANCE TO Sphenophorus venatus vestitus Billbug ( Sphenophorus spp.) damage in turfgrass is often misdiagnosed as drought-stress or disease (Niemczyk and Shetla r 2000), leading to inappropriat e management efforts. Early instar larvae are stem borers, late instar larvae are root feeders, and adults feed on stem tissue. However, the late instar larvae are considered the most damaging stage (Satterthwait 1931). The most abundant and damaging billbug species in warm season turfgrass is the hunting billbug, S. venatus vestitus. Sphenophorus venatus vestitus has multiple generations with overlapping life stages each year in Florida (Chapt er 2), which makes infestations of S. venatus vestitus difficult to consistently manage with either preventive or curative insecticides. Its primary hosts are bermudagrass (Cynodon dactylon (L.) Pers.) and zoysiagrass ( Zoysia spp.). An alternative, non-chemical control strategy is to develop cultivars that are more resistant to billbug feeding damage. Resistance in differe nt bermudagrass cultivars has been identified against key arthropod pests (Qui senberry 1990) such as mites (Tashiro 1987), fall armyworm ( Spodoptera frugiperda J.E. Smith) (Quisenberry and Wilson 1985, Jamjanya and Quisenberry 1988, Croughan and Quisenberry 1989), and mole crickets ( Scapteriscus spp.) (Reinert and Busey 1984). Both non-preference an d antibiosis to fall armyworm were identified in different zoysiagrass cultivars (Chang et al. 1985). A host range studies on S. parvulus was done by Kindler et al. (1983), they f ound only Kentucky blue grass, Poa pratensis L. showed significant infestation of S. parvulus larvae. Bermudagrass cultivars such as Tifway, Tifdwarf and Tifgreen have been identified as having high mite, Eriophyes cynodoniensis Sayed, resistance (Tashiro 1987); Zoysia tenuifolia was also identified highly re sistant to the Banks grass mite, Oligonychus pratensis Banks (Busey 1982).

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43 However, the first report of zoys iagrass cultivar resistance to S. venatus vestitus was by Reinert (2001). The objective of this study was to document the impact to feeding damage by S. venatus vestitus on a range of bermudagrass a nd zoysiagrass cultivars. Materials and Methods Damage potential of adult S. venatus vestitus on Fur Brmudagrass Cltivars Plant material Ten plugs (1 0.2 cm diameter) of each of the bermudagrass cultivars Celebration, Tifdwarf, Tifeagle, and Tifway we re obtained from established plot s at the University of Florida Plant Science Unit in Citra, FL, on 30 March 2007. Plugs were allowed to become established in pots (11.4 cm diameter) with native soil (sandy lo am) in a greenhouse. Pots were fertilized with 113.5 g of Miracle-Gro all purpose fertilizer (20-20-20) each week and irrigated daily. Adult damage potential Adult S. venatus vestitus were collected from linear pitf all traps at Gainesville Country Club, FL, on 15 May 2007. Ten adult males or female s of unknown age were placed onto the pot of each bermudagrass cultivar (five pots per cultivar), and confined with a fine white mesh. After 2 weeks, pots were destructively sampled, adult survival per pot was determined, and the location of male or female notching damage, numb er of notches per pot, notch length and width, and diameter of damaged area were measured unde r a 10X dissecting microscope with a caliper. The average daily temperature in the greenhous e was 16-30C, average soil temperature was 19.4C, light intensity was 9,688 lum/m2 with 14:10 hr (L:D). Data were analyzed using a twoway ANOVA (SAS Institute 2000) to detect the effect of cultivar and billbug sex on adult S. venatus vestitus damage potential. Means were co mpared using Tukeys HSD test ( P < 0.05).

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44 Bermudagrass Cltivar Peference Test Plant material Cores of bermudagrass cultiv ars Celebrati on, Tifdwarf, Tifeagle, and Tifway were obtained from the University of Florida Plan t Science Unit in Citra, FL, on 24 August 2007. Cores were planted into 8.9 cm di ameter pots with native soil, fe rtilized weekly with 113.5 g of Miracle-Gro all purpose fertilizer (20-20-20) and ir rigated daily. Grass height was maintained at about 5 cm. The test arena (22 22 10 cm) had fourteen holes (6 mm diameter) in the bottom to allow drainage, and an identical cont ainer with its bottom removed was glued upside down on top of it (Figure 3-1). One pot of each cultivar was randomly placed in the container, and the area between pots was fille d with potting soil (Scotts Co. Ge rvais, OR) so the soil was 10 cm deep within and between pots. Preference evaluation Four of each unknown-aged adult m ale and female S. venatus vestitus were collected from the U. F. Plant Science Unit in Citra, FL, on 20 November 2007 and released in the center of each container between the pots within 24 hours. All adults were removed from containers 30 days after introduction (19 D ecember 2007). All pots were evaluated on 9 10 January 2008 for the total number of eggs and la rvae in the stems or soil, and the total number of adult feeding notches. Data were analyzed using a one-way ANOV A (SAS Institute 2000) to detect the effect of variety on adult S. venatus vestitus preference. Means were comp ared using Tukeys HSD test ( P < 0.05). Zoysiagrass Cultivar Resistance to S. venatus vestitus Plant material Pots (7.6 cm diam eter) of 17 zoysiagrass cu ltivars were established with sand in a greenhouse in Gainesville, FL, for 90 days. Cultiv ars included Belair, Cashmere, Cavalier,

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45 Compadre, Crowne, Diamond, El Toro, E merald, Empire, J amur, Palisades, Pristine Flora, Royal, Ultimate Flora, Zenith, Zeon and Zorro. Pots were fertilized with 113.5 g of Miracle-Gro all purpose fertilizer (20-20-20) per week, irrigated as needed, and turf was maintained at ~ 3.8 cm height. Resistance evaluation Adult S. venatus vestitus were collected from the pitfall traps at LaGorce Country Club in Miami, FL, on 28 December 2006 and transported to the laboratory in Gainesville. Two males and two females were placed onto each pot within 48 hours (five pots per cultivar), and confined with a fine white mesh. Four weeks after adult caging, pots of each cultivar were rated for their percentage of live cover (0-100%), color (1 = brown, 9 = dark green), density (1 = thin, 9 = dense), overall quality (1 = poor, 9 = excellent), and the percentage of turf infested with the armored scale Duplachionaspis divergens Green. Pots were also destructively sampled to recover all life stages of S. venatus vestitus. Data were analyzed using a one-way ANOVA (SAS Institute 2000) to detect the effect of cultivar difference on adult S. venatus vestitus feeding potential and oviposition. Means were separate d using Tukeys HSD test ( P < 0.05). This test was repeated with 18 cultivars (Meyer was added). Ten pots (7.6 cm diameter) per cultivar were planted in sand in Novemb er 2007, allowed to establish until January 2008, and maintained as previously described. Adult S. venatus vestitus were collected from the U.F. Plant Science Unit in Citra, FL, on 11 January 2008, a nd two pairs of males and females were placed onto each pot within 24 hours. Each pot was encirc led with clear plastic (15 cm high) to confine adults to the pot and allow normal light intensit y and air flow. Four week s after caging, pots of each cultivar were rated for thei r percentage of live cover, color, density, overall quality, and the percentage of turf infested with D. divergens Pots were also destructiv ely sampled to recover all

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46 life stages of S. venatus vestitus. Data were analyzed using a one-way ANOVA (SAS Institute 2000) to detect the effect of cultivar difference on adult S. venatus vestitus feeding potential and oviposition. Means were separate d using Tukeys HSD test ( P < 0.05). Results and Discussion Damage Potential of Adult S. venatus vestitus on Four Be rmudagrass Cultivars Both male and female S. venatus vestitus caused damage, presumably by feeding, to the bermudagrass stolons and rhizomes. The notch es were 3.6 0.3 mm below the nearest crown for males and 4.0 0.2 mm below for females in a ll varieties (Table 3-2). Notch length was similar for both sexes (2.3 0.2 mm for males, 2.4 0.1 mm for females), and notch width was identical (0.7 0.03 mm). Females made more notches in pots of Tifway than in other cultivars ( F = 3.83; df = 3, 19; P = 0.0305), but males made a similar numb er of notches in pots of each cultivar (Table 3-1). Eggs were found in notch es on stems or leaf sheath about 1.3 0.1 mm wide. The diameter of the damaged area was signif icantly thicker in Tifway and Celebration than in Tifeagle and Tifdwarf ( F = 8.2; df = 3, 39; P = 0.0003). It is possible that billbugs may preferentially attack cultivars with thicker st em diameters, such as Tifway and Celebration compared to Tifeagle and Tifdwarf (Kenwo rthy, 2008 personal communication). Thicker stems may indicate greater plant resources for their o ffspring. Turfgrass thatch thickness and density may also influence billbug populations by providing greater humidty and shade. For example, populations of S. parvulus were larger in areas with thicke r thatch(Kindler and Spomer 1986). Bermudagrass Cultivar Preference Test One m onth after adult billbugs were caged onto pots, zero to two larvae and 0-25 notches occurred in pots of each cultivar. Tifway bermudagrass had the most of larvae and notches recorded, and Tifeagle had the least (Table 3-2). The mean number of larvae was significantly higher in Tifw ay than in Tifeagle ( F = 3.49; df = 3, 39; P = 0.025). The mean

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47 number of notches from adult feeding was signi ficantly higher in Tifway than the other three cultivars (F = 15.62; df = 3, 39; P < 0.0001). Sphenophorus ventaus vestitus fed more, layed more eggs, and caused more damage on Tifway bermudagrass than on the other bermudagra ss cultivars tested. This may either indicate greater preference for this cultiv ar or compensatory feeding. Fu rther study to determine the host quality and the impact on offspring survival, development time and fecundity are needed. Currently there is no resear ch documenting bermudagrass cultivars having resistance to S. ventaus vestitus feeding. The genetic resistance among bermudagrass cultivars is unknown, however, a broad range of gr ass species resistant to S. parvulus based on genetic differences was identified (Asay et al. 1983). But they only rated the damage visually on each grass trial in the field without recording number of notches on stem tissue or larv ae present in the soil. Visual damage may not have been the result of billbugs. Zoysiagrass Cultivar Resistance to S. venatus vestitus For the first run of this experim ent Diamond had the best performance four weeks after adult billbug infestation fo r percent living coverage ( F = 6.16; df = 16, 254; P < 0.0001), color ( F = 4.36; df = 16, 254; P < 0.0001), density ( F = 17.48; df = 16, 84; P < 0.0001) and quality ( F = 7.63; df = 16, 254; P < 0.0001). Belair performed the poorest for percent living coverage (F = 6.16; df = 16, 254; P < 0.0001) and color ( F = 4.36; df = 16, 254; P < 0.0001). Compadre and Empire had the least dense ( F = 17.48; df = 16, 84; P < 0.0001). Belair and Compadre had the poorest turf quality ( F = 7.63; df = 16, 254; P < 0.0001) (Table 3-3). Compadre, Diamond, Emerald, and Royal were more susceptible to the scale, D divergens ( F = 14.98; df = 16, 254; P < 0.0001). Notches made by adults were 0.8 to 3.9 mm long and 0.4 t o1.0 mm wide across cultivars (Table 3-4).

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48 For all cultivars, zero to eight eggs + larvae and one to thir teen notches were found in pots 4 weeks after adults were caged (Table 3-4). Th e mean number of notches and the mean number of eggs + larvae were not significantly different among two zoysiagrass species or the hybrids. Regardless of species, the mean number of eggs and larvae combined was significantly higher in Crowne, El Toro, and Jam ur than other cultivars ( F = 1.91; df = 16, 84; P = 0.0338), but the mean number of notches was not signi ficantly different among cultivars. After evaluating the second test, 0-12 eggs and larvar, 4-38 notches were found per pot (Table 3-5). Among Z. japonica cultivars, the mean number of notches, and the mean number of eggs + larvae combined were not significantly different. Among Z. matrella cultivars, the mean number of notches ( F = 4.52; df = 5, 29; P = 0.0048), and eggs plus larvae combined were significantly higher in Cashmer e than other cultivars ( F = 3.5; df = 5, 29; P = 0.0162). Among hybrid cultivars, the mean number of notches, and the mean number of eggs + larvae combined were not significantly different. Regardless of cultivar, the mean number of notches, and the mean number of eggs and larvae combined was significantly higher in Z. japonica than other species (F = 24.02; df = 2, 89; P < 0.0001) ( F = 14.5; df = 2, 89; P < 0.0001), respectively. Regardless species, significantly more notches oc curred in Meyer, Zenith, Compadre, and Palisades than in other cultivars ( F = 5.01; df = 17, 89; P < 0.0001). The mean number of eggs and larvae combined was significantly higher in Meyer, Crowne, Compadre, Zenith, and El Toro than other cultivars ( F = 2.81; df = 17, 89; P = 0.0012). Reinert (2001) reported that four cultivars of Z. matrella (Diamond, DALZ 9601, Cavalier and Royal) are resistant to S. venatus vestitus, and Meyer is the most susceptible among nine cultivars. In this study, cultivars with the highest number of eggs + larvae such as Compadre, Crowne, Jamur, Meyer, and Zenith were all in Z. japonica Z. matrella cultivars

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49 generally produced fewer S. venatus vestitus eggs or larvae. with less offspring. Total number of damage was also higher in Z. japonica cultivars than in Z. matrella cultivars. However, billbugs feeding more on cultivars of Z. japonica did not mean they were more susceptible since I did not evaluate the nutrient leve l of each cultivar. Thus this feedi ng behavior could be compensatory. These findings plus the rating data indicate that cultivars of Z. matrella have better resistance against S. venatus vestitus feeding and oviposition. The reason I found mostly larvae in the first ev aluation and mostly eggs in the second time evaluation remains unknown. I specula te that the warmer temperat ure of the greenhouse in the first time evaluation speed up egg hatch to larvae. In addition, geneti c resistance in grass cultivars to insect pest may be influenced by so il fertility and cultural conditions in which the grass is grown (Quisenberry 1990). Therefore, to eliminate all the potentia l factors that might affect the results, such as grass resources, pot size, amount of fertilizer I did both bermudagrass and zoysiagrass trials in our greenhouse rather than in the field. However, two different populations of S. ventaus vestitus were used from different locatio ns (Gainesville and Miami) in the test, and were used in different seasons, which may have influenced test results. To conclude, cultivars of both bermudagrass and zoysiagrass appear to have varying levels of susceptibility to S. venatus vestitus Turfgrass managers may be able to reduce their pesticide use by selecting less susceptible cultivars when installing, renovating, or repairing turfgrass areas.

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50 Table 3-1. Damage between male and female S. venatus vestitus on four bermudagrass cultivars in the greenhouse. Cultivar Billbug gender Mean no. notches (SEM)1/pot Notch length (mm) Notch width (mm) Diameter of damaged area (mm) Celebration Male 21.2 2.1 2.0 0.2 0.7 0.04 1.5 0.1 Tifdwarf Male 29.6 3.7 1.8 0.1 0.8 0.03 0.9 0.04 Tifeagle Male 24.8 3.2 1.8 0.1 0.6 0.02 0.7 0.02 Tifway Male 22.0 4.8 2.3 0.2 0.7 0.04 1.2 0.04 Celebration Female 23.2 2.1b 2.4 0.2 0.8 0.04 1.3 0.1 Tifdwarf Female 26.8 1.8b 1.7 0.1 0.8 0.03 1.1 0.03 Tifeagle Female 26.2 2.8b 2.0 0.1 0.6 0.03 0.7 0.02 Tifway Female 34.8 3.1a 2.4 0.1 0.7 0.03 1.3 0.04 1 Means within columns with different letters are statistically different at = 0.05 ( F = 3.83; df = 3, 19; P = 0.0305)

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51 Table 3-2. Mean (SEM) number of notches and surviving larvae per pot on four bermudagrass cultivars in the greenhou se preference test. Cultivar Mean1 no. larvae/pot Mean2 no. notches /pot Celebration 0.4 0.2b 6.3 0.9b Tifdwarf 0.3 0.2b 6.1 2.0b Tifeagle 0b 2 0.4b Tifway 0.7 0.2a 15.7 1.9a 1 Means within columns with different letters are statistically different at = 0.05 ( F = 3.49; df = 3, 39; P = 0.025) 2 Means within columns with different letters are statistically different at = 0.05 ( F = 15.62; df = 3, 39; P < 0.0001)

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52 Table 3-3. Ratings (SEM) of seventeen zoys iagrass cultivars after 4 weeks of adult S. venatus vestitus infestation. Species/Cultivar Percent live cover 1Color (1-9)2 Density (1-9)3Quality (1-9)4 % D. divergens5 Zoysia japonica Belair 34 4.3e 4.5 0.4d 4.2 0.4cde 2.9 0.2e 10.3 2.9c Compadre 50 3.1bcde 6.1 0.4abcd3.3 0.3e 2.9 0.2e 45.3 8.3a Crowne 60.7 3.3abcd 6.1 0.3abcd5.5 0.3bcd 4.6 0.4bcde 2.0 0.8c El Toro 41.7 7.8de 5.8 0.1abcd6.7 0.3ab 3.8 0.6cde 4.3 1.0c Empire 60.0 4.7abcd 6.4 0.3abc 3.3 0.3e 3.2 0.3de 18.3 3.8bc Jamur 66.0 3.1abc 6.3 0.4abc 5.5 0.4bcd 5.0 0.4abcd 9.7 2.6c Palisades 68.7 4.6ab 6.5 0.3ab 4.2 0.3cde 4.6 0.3bcde 6.3 1.4c Ultimate Flora 57.3 3.5abcd 6.5 0.4ab 5.5 0.4bcd 4.2 0.4bcde 7.3 1.3c Zenith 43.3 4.9cde 4.9 0.4bcd 4.1 0.4de 3.6 0.5de 15.0 5.4c Zoysia matralla Cashmere 58.7 3.6abcd 5.4 0.3abcd5.9 0.3bc 4.9 0.2abcd 2.3 0.7c Cavalier 47.7 6.9bcde 4.8 0.7cd 6.9 0.3ab 3.7 0.6cde 19.7 6.3bc Diamond 79.6 3.6a 6.9 0.3a 7.7 0.4a 6.7 0.3a 40.7 6.5ab Pristine Flora 63.3 3.6abcd 5.9 0.3abcd6.2 0.4ab 5.1 0.3abcd 5.3 1.4c Royal 70.0 4.6ab 6.8 0.2a 7.1 0.3ab 5.7 0.3ab 54.3 8.2a Zeon 56.7 6.0bcde 6.1 0.3abcd7.6 0.3a 5.5 0.4abc 11.7 3.8c Zorro 58.3 4.2abcd 6.3 0.2abc 7.0 0.3ab 5.0 0.4abcd 10.3 3.0c Hybrid Emerald 50.0 3.5bcde 6.6 0.3a 5.5 0.4bcd 3.9 0.3bcde 50.7 6.9a 1 Means within columns with different letters are statistically different at = 0.05 ( F = 6.16; df = 16, 254; P < 0.0001). 2 Means within columns with different letters are statistically different at = 0.05 ( F = 4.36; df = 16, 254; P < 0.0001). 3 Means within columns with different letters are statistically different at = 0.05 ( F = 17.48; df = 16, 84; P < 0.0001). 4 Means within columns with different letters are statistically different at = 0.05 ( F = 7.63; df = 16, 254; P < 0.0001). 5 Means within columns with different letters are statistically different at = 0.05 ( F = 14.98; df = 16, 254; P < 0.0001)

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53 Table 3-4. Mean number (SEM) of S. venatus vestitus eggs and larvae, and characteristics of adult notches on 17 zoysiagrass cultivars, after 1 month of adult confinement. Species/Cultivar # Mean no. eggs and larvae/pot Mean no. notches /pot Notch Length (mm) Notch Width (mm) Zoysia japonica Belair 0 1.4 0.5 3.1 0.7 0.6 0.1 Compatibility 0 1.0 0.4 1.5 0.4 0.6 0.1 Crowne 1.6 0.7 2.6 1.1 1.7 0.4 0.6 0.1 ElToro 1.2 0.6 1.6 0.7 1.3 0.2 1.0 0.1 Empire 0.8 0.5 2.0 0.9 1.2 0.3 0.6 0.1 Jamur 1.4 0.9 1.6 1.2 3.6 0.7 1.0 0.2 Palisades 0 0.6 0.2 1.1 0.2 0.5 0.03 Utimate Flora 0.4 0.4 1.2 0.7 1.3 0.2 0.7 0.1 Zenith 0 0.2 0.2 0.8 0.4 Zoysia matrella Cashmere 0.8 0.6 2.4 1.2 3.3 0.9 0.5 0.1 Cavalier 0.4 0.2 1.2 0.8 1.4 0.2 0.5 0.04 Diamond 0 0.6 0.4 0.8 0.1 0.4 0.03 Pristine Flora 0 1.4 0.7 3.6 0.9 0.6 0.1 Royal 0.2 0.2 2.2 1.1 2.6 1.1 0.6 0.1 Zeon 0 1.2 0.4 1.6 0.4 0.5 0.02 Zorro 0.2 0.2 1.0 0.3 3.9 1.2 0.6 0.1 Hybrid Emerald 0 0.4 0.4 1.4 0.8 0.4 0.1

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54 Table 3-5. Mean number (SEM) of S. venatus vestitus eggs and larvae, and adult notches on 18 zoysiagrass cultivars, after one month of adult confinement. Species/Cultivar Mean no. eggs and larvae/potMean1 notches/pot Zoysia japonica Belair 0.8 0.4 5.6 1.9abcd Compadre 2.4 0.9 6.2 1.5abc Crowne 2.6 0.7 3.2 0.9abcd ElToro 1.8 0.6 3.4 0.7abcd Empire 0.8 0.4 5.6 1.9abcd Jamur 1.2 0.5 4.8 0.9abcd Meyer 3.0 0.9 6.8 1.2ab Palisades 1.0 0.5 5.8 0.7abcd Utimate Flora 0.2 0.2 0.8 0.4 Zenith 2.6 1.7 7.6 1.3a Zoysia matrella Cashmere 1.0 0.3 5.4 1.1abcd Cavalier 0.4 0.2 1.4 0.2cd Diamond 0 1.6 0.8bcd Pristine Flora 0 1.8 0.4bcd Royal 0.2 0.2 1.6 0.5bcd Zeon 0.2 0.2 1.0 0.4cd Zorro 0 2.6 1.0abcd Hybrid Emerald 0 1.0 0.3cd 1 Means within columns with different letters are statistically different at = 0.05 ( F = 5.01; df = 17, 89; P < 0.0001)

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55 Figure 3-1. Sphenophorus venatus vestitus preference test for bermudagrass cultivars.

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56 CHAPTER 4 EFFECT OF ENDOPHYTE LEVEL IN PERE NNIAL R YEGRASS ON THE SURVIVAL AND DEVELOPMENT OF Sphenophorus venatus vestitus Endophytic fungi and phytophagous insects acquire nutrients from sharing the same host plant and interacting with one another (Wilson a nd Carroll 1997). Studies have documented that the endophytic fungus Neotyphodium lolii Latch associated with gr asses such as perennial ryegrass ( Lolium perenne L.) or tall fescue (Festuca arundinacea Schreb) can have enhanced resistance to herbivory through the production of alkaloids (Prestidge and Gallagher 1988, Kunkel et al. 2003). Some insects can detect en dophyte infected grass and avoid feeding on it, while those that do feed on it suffer reduced f itness, survival and f ecundity (Johnson-Cicalese 1997). Endophytes occur in 13 genera of grasses including bluegra ss, bentgrass, fescue, and ryegrass, and affect 40 insect species from six different orders (Johnson-Cicalese 1997). Endophytic tall fescue was more e ffective against first and second instar white grubs than third instars (Potter et al. 1992, Koppenhfer et al. 2003). Ahmad et al. (1986) reported that a reduction of bluegrass billbug ( Sphenophrous parvulus Gyllenhal) larval population density and feeding damage was associated with endophytic perennial ryegrass. Subsequently, Richmond et al. (2000) reported that, in general, visu al damage and the larval numbers of S. parvulus decreased as the proportion of endophytic perennial ryegrass incr eased. Another field trial in New Jersey indicated that endophy tic tall fescue provided a high level of resistance to the damage of four billbug species (Murphy et al. 1993). Endophyte-en hanced turfgrass resistance also affects populations of sod webworm (Murphy et al. 1993 ), Argentine stem weevil, Listronotus bonariensis (Kuschel) (Prestidge 1988) and gall-forming insects, Besbicus mirabilis Kinsey (Wilson and Carroll 1997). However, this is the first report on th e effect of pure and overseeded endophytic perennial ryegrass on the hunting billbug (S. venatus vestitus Chittenden).

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57 Materials and Methods Sphenophorus venatus vestitus Survival, Development, and Da mage on Four Endoyphytic Perennial R yegrass Cultivars To assess the direct effect of endophytic perennial ryegrass on the survival and development of S. venatus vestitus, a greenhouse experiment wa s conducted. The cultivars Citation Fore (76% endophyte), Cat alina II (72%), Bri ghtstar SLT (80%), and Caparral II (6%) (Scotts Company) were seed ed at a rate of 29.4 kg / 1,000 m2 with 80% sand mixed with 20% top soil in Plastic pots (11.7 cm diameter) on 17 November 2007. The endophyte level of each cultivar was evaluated by the Agrinostics Company. Each treatment was replicated five times in a randomized complete block design. Miracle-Gro all purpose fertilizer (113.5 g, 2020-20) was applied to each pot weekly and pots were irrigated daily. Each pot was infested with two male and two female adult S. venatus vestitus collected from Tifway bermudagrass at the U.F. Plant Science Research Unit in Citra, FL, on 10 January 2008. Pots were encircled up to 15 cm above the turf height with a clear plastic tube to preven t adult escape. Turfgrass damage was visually rated (1 = little damage 9 = severe damage), and adult survival, and the number of eggs and larvae found in stem tissue or soil were dete rmined after 2 weeks (24 January 2008). Data were analyzed using a one-way ANOVA (SAS Institute 2000) to det ect the effect of different cultivars on adult S. venatus vestitus damage potential and oviposition. Means were compared using Tukeys HSD test ( P < 0.05). Impact of Overseeding Two Bermudagrass Cult ivars w ith Endophytic Perennial Ryegrass on S. venatus vestitus Survi val, Development, and Damage To assess the effect of overseeding two berm udagrass cultivars with a perennial ryegrass cultivar with a high percentage of en dophyte on the survival and development of S. venatus vestitus a greenhouse experiment was conducted. Co res (10.2 cm diameter) of Tifway and Tifeagle bermudagrass were obtained from the U. F. Plant Science Research Unit in Citra, FL,

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58 and planted into 11.7 cm diameter plastic pots on 27 April 2007, and were allowed to establish with native soil (sandy loam). Five pots of each bermudagrass cultivar were overseeded on 17 November 2007 at a rate of 50.5 kg / 1,000 m2 with Citation Fore, a perennial ryegrass cultivar with 76% endophyte. Five additional pots of each bermudagrass cultivar not overseeded. Miracle-Gro all purpose ferti lizer (113.5 g, 20-20-20) was applied to each pot weekly and pots were irrigated daily. Each pot was infest ed with two adult male and two female S. venatus vestitus collected from the U.F. Plant Science Re search Unit in Citra, FL, on 10 January 2008. Each pot was encircled up to 15 cm above the turf height with a clear plastic tube to prevent adult escape. Pots were destructively evaluated and th e total number of eggs and larvae found in stem tissues and soil, and number of adult not ches made, were recorded after 2 weeks (28 January 2008). Data were analyzed using a twoway ANOVA (SAS Institute 2000) to detect the effect of different cultiv ars and overseeding on adult S. venatus vestitus damage potential and oviposition. Means were compared using Tukeys HSD test ( P < 0.05). Overseeded Bermudagrass Field Trial Due to lim itations in seed availability, two di fferent perennial ryegrass cultivars were used in the field study. A cultivar with a low endophyte level (SR4500, 33% endophyte) and a cultivar with a high endophyte level (SR4420, 78% endo phyte) were overseeded onto plots (1.83 1.83 m) of Tifway bermudagrass at the U.F. Plan t Science Research Unit in Citra, FL, on 9 November 2006 at a rate of 50.4 kg/1,000 m2 using a Scotts Proturf professional drop spreader. Control plots were not overseeded. Bermudagra ss height was ~1 cm and thatch was ~1.4 mm thick. Each ryegrass treatment was replicated te n times, but there was only space for five control plots (25 total plots), in an unbalanced randomized complete block design. About 950 S. venatus vestitus and 450 S. inaequalis adults were collected from Gainesville golf courses and released onto the plots to establish a population during la te summer and fall of 2006, before overseeding

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59 occurred. Turf and soil cores (10.2 cm diameter, 8 cm depth) were removed from each plot on 19 November 2006 (3 cores per plot), 20 January (2 cores per plot) and 2 April 2007 (5 cores per plot) and destructively examined for all billbug life stages. Data were analyzed with two-way ANOVAs (SAS Institute 2000), and if significant, means were separated by Tukeys HSD test (SAS Institute 2000). Results and Discussion Sphenophorus venatus vestitus Survival, Development, and Da mage on Four Endoyphytic Perennial R yegrass Cultivars No eggs, larvae, or adult notching damage were found in the pots of any of the four ryegrass cultivars after 2 weeks of being infested with adult S. venatus vestitus, regardless of endophyte level. The adult survival of each cultivar was 95% in Citation Fore, 80% in Catalina II, 85% in Brightstar SLT, and 95% in Caparral II. Females were not dissected to examine the number of eggs in their ovaries af ter 2 weeks of caging. Only some grass blades turned yellowish and shrank, especi ally in Caparral II. Perennial ryegrass is considered a host for S. venatus vestitus (Jonhson-Cicalese and Funk 1990), although I did not measure the stem diameter, I assumed perhaps the stem systems we re not developed or th ick enough to allow first instar feeding, so oviposition by female S. venatus vestitus had not occurred. Given the lack of notching damage, adults may have been repelled or not induced to feed and oviposit. Ants and spiders can detect alkaloids from their poten tial preys integument (Montllor et al. 1991, Schaffner et al. 1994), so perhaps S. venatus vestitus can also sense the presence of alkaloids from endophyte-infected turfgrass cultivars. Impact of Overseeding Two Bermudagrass Cult ivars w ith Endophytic Perennial Ryegrass on S. venatus vestitus Survival, Development, and Damage This is the first study to examine the e ffect of overseeding with endophyte-enhanced perennial ryegrass on S. venatus vestitus for potential use as a management tool in the southern

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60 United States. However, Murphy et al. (1993) demonstrated that endophyte-e nhanced tall fescue ( Festuca arundinacea Schreb) reduces the feeding damage of S. venatus vestitus and S. minimus. From all ryegrass cultivars in th is study, zero to three larvae, and three to 40 notches were found in pots after 2 weeks of being in fested with adults. Eggs were not found in all pots. The mean number of notches and larvae we re significantly higher in non-ove rseeded Tifway pots than in other cultivars ( F = 32.44; df = 3, 19; P < 0.0001) and (F = 10.43; df = 3, 19; P = 0.0005), respectively. Tifeagle overseed ed with endophytic perennial ryeg rass had the least damage and no larvae were found (Table 4-1). Regardless of cultivar, the mean number of notches and larvae were significantly higher in contro l pots than in overseeded pots ( F = 25.41; df = 1, 19; P < 0.0001) and ( F = 18.78; df = 1, 19; P = 0.0004), respectively. It is unclear whether adults avoided feeding and ovipositing on overseeded bermudagrass, or if eggs and early instar larvae died in the overseeded pots. Similarly, in th e northern U.S., overseed ing Kentucky bluegrass ( Poa pratensis L.) with endophytic perennial ryegrass can reduce S. parvulus larval populations and damage (Richmond et al. 2000). Overseeded Bermudagrass Field Trial Despite releasing over 1,000 Sphenophorus spp. adults into the be rm udagrass plots before overseeding was done, only one firs t instar was found in a stolon from a control plot in the November 2006 sample, one mature larva was found in each of two control plots and one in an SR 4500 (low endophyte) plot in the January 2007 sample, and one mature larva was found in a control plot and one from SR 4500 in the April 2007 sample. Grass roots and stems were healthy in all turf cores; no obvious da mage or notches were visible. The number of billbugs collected from these samples was too low to detect treatme nt differences. It is pos sible that a large enough billbug population did not develop on the plots, an d that the number of cores taken at each sampling date was insufficient. However, results from caging only two pairs of S. venatus

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61 vestitus in greenhouse pot trials demons trates that adults can cause significant turfgrass damage and lay several eggs in small areas. The fi eld plots were surrounded by other potential nonoverseeded turfgrass areas, so it is also possible that adults moved out of the desired area (they were not confined with cages). Other overseeding field studies were done in areas with a history of billbug infestation, rather than newly infested areas (Murphy et al. 1993, and Richmond et al. 2000). Turfgrass height could also affect female billbug oviposition choice and/or larval survival, if it impacts stem or root diameter and density.

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62 Table 4-1. The effect of overseeding with endoph ytic perennial ryegrass on two bermudagrass cultivars on S. venatus vestitus damage potential and oviposition Cultivar Treatment Mean no. notches/pot (SEM)1 Mean no. larvae/pot (SEM)2 Tifway Control 31.2 3.3a 1.8 0.4a Tifeagle Control 18.4 1.2b 0.8 0.4b Tifway Overseeded 13.2 1.0b 0b Tifeagle Overseeded 5.6 0.9c 0b 1 Means within columns with different letters are statistically different at = 0.05 ( F = 32.44; df = 3, 19; P < 0.0001). 2 Means within columns with different letters are statistically different at = 0.05 ( F = 10.43; df = 3, 19; P = 0.0005).

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63 APPENDIX A EVALUATION OF BEMUDAGRASS RESISTANCE TO Sphenophorus inaequalis Materials and Methods Damage Potential of Adult S. inaequalis on Four Bermudagrass Cultivars Plant material Ten plugs (1 0.2 cm diameter) of each of the bermudagrass cultivars Celebration, Tifdwarf, Tifeagle, and Tifway were obtaine d from established plots at the University of Florida Plant Science Unit in Citra, FL, on 30 May 2006. Plugs were allowed to become established in pots (11.4 cm diam eter) with native soil (sandy loam) in a greenhouse. Pots were fertilized with 113.5 g of Miracle-Gro all purpose fertilizer (20-20-20) each week and irrigated daily. Adult damage potential Adult S. inaequalis were collected from linear pitfall traps at West End Country Club in Gainesville, FL, on 8 August 2006. Ten unaged adult ma les or females were placed onto the pot of each bermudagrass cultivar (five pots per cultivar), and confined with a fine white mesh. After 2 weeks, pots were destructively sampled, adult survival per pot was determined, and the location of male or female notching damage, numb er of notches per pot, notch length and width were recorded. In the S. inaequalis test the average daily temperature in the greenhouse was 2132C, average soil temperature was 21C, light intensity was 10,764 lum/m2 with 14:10 hr (L:D). Data were analyzed using a two-way ANOVA (SAS In stitute 2000) to detect the effect of variety and billbug sex on adult damage potential. Mean s were compared using Tukeys HSD test ( P < 0.05).

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64 Results and Discussion Damage Potential of Adult S. inaequalis on Four Bermudagrass Cultivars The notches caused by S. inaequalis were 3.0 0.3 mm below crown for males and 4.1 0.3 mm below for females in all varieties (Tab le 3-1). Average notch length was 2.0 0.2 mm for males and 1.9 0.1 mm for females. Averag e notch width was 0.7 0.04 mm for males and 0.7 0.03 mm for females. The mean number of not ches from both male and female feeding was significantly higher in Tifdwarf than other cultivars (males: F = 35.1; df = 3, 19; P < 0.05; females: F = 16.1; df = 3, 19; P < 0.05). The mean number of notches caused by females feeding across cultivars was significantly higher than males ( F = 5.81; df = 1, 39; P < 0.05) (Table 3-1). In the bermudagrass variety trial, S. inaequalis tended to feed more on Tifdwarf bermudagrass than other cultivars, and female S. inaequalis also fed more than males under a nochoice environment. The reason why Tifdwarf was the most susceptible variety to S. inaequalis feeding among the four bermudagrass varieties is not clear understood. One reason might be the less resistance from the cultivar itself. Tashir o (1987) reported a high m ite resistance in the bermudagrass cultivars Tifdwarf and Tifway. Howeve r, Tifdwarf apparently has less resistance against to S. inaequalis feeding comparing to other cultivars. It is reasonable that females cause more notches than males because the needs of energy for ovary development and oviposition.

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65 Table A-1. Damage between male and female S. inaequalis on four bermudagrass cultivars in the greenhouse Cultivar Billbug gender Mean1, 2 no. notches (SEM)/pot Notch length (mm) Notch width (mm) Diameter of damaged area (mm) % adult survival Celebration Male 16.2 1.5b 1.7 0.1 0.6 0.02 1.0 0.04 70 Tifdwarf Male 34.6 2.0a 1.6 0.1 0.6 0.02 0.7 0.02 86 Tifeagle Male 18.6 1.7b 1.0 0.1 0.4 0.01 0.7 0.02 84 Tifway Male 12.2 1.4b 2.0 0.2 0.7 0.04 1.2 0.06 64 Celebration Female 20.8 1.5b 1.7 0.1 0.7 0.03 1.1 0.04 70 Tifdwarf Female 42.4 3.2a 2.5 0.2 0.8 0.03 1.0 0.03 82.5 Tifeagle Female 20.4 2.1b 1.0 0.1 0.4 0.01 0.7 0.02 76 Tifway Female 28.8 3.1b 1.9 0.1 0.7 0.03 1.2 0.03 82 1 Means within columns with different letters are statistically different at = 0.05 ( F = 35.1; df = 3, 19; P < 0.05). 2 Means within columns with different letters are statistically different at = 0.05 ( F = 16.1; df = 3, 19; P < 0.05).

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66 LIST OF REFERENCES Ahmad, S., J. M. Johnson-Cicalese, W. K. Dickson, and C. R. Funk. 1986. Endophyteenhanced resistance in perennial ryegrass to the bluegrass billbug, Sphenophorus parvulus Entom ol. Expt. Appl. 41: 3-10. Asay, K. H., and J. D. Hansen, B. A. Haws, and P. O. Currie. 1983. Genetic differences in resistance of range grasse s to the bluegrass billbug, Sphenophorus parvulus (Coleoptera: Curculionidae). J. Range Management. 36: 771-772. Brandenburg, R. L. and M. G. Villani. 1995. Handbook of Turfgrass Insect Pests. Entomol. Soc. Am. Lanham, MD. Buss, E. A. 2003. Insect pest management on golf courses. http://edis.ifas.ufl.edu/IN410 UF/IFAS Extension. Gainesvi lle. Florida. Com plete. Buss, E. A. and J. B. Unruh. 2006. Insect management in your Florida lawn. http://edis.ifas.ufl.edu/LH034 UF/IFAS Extension. Gainesville. Florida. Com plete. Buss, E. A., and J. C. Turner. 2004. Insect pest management on turfgrass. http://edis.ifas.ufl.edu/IG001 UF/IFAS Extension. Gainesville. Florida. Com plete. Buss, E. A., J. L. Capinera, and N. C. Leppla. 2006. Pest mole cricket management. http://edis.ifas.ufl.edu/LH039 UF/IFAS Extension. Gainesville. Florida. Com plete. Chang, N. T., B. R. Wiseman, R. E. Lynch, and D. H. Habeck. 1985. Fall armyworm: expressions of antibiosis in selected grasses. J. Entomol. Sci. 20: 179-188. Croughan, S. S., and S. S. Quisenberry. 1989. Enhancement of fall armyworm (Lepidoptera: Noctuidae) resistance in bermudagrass th rough cell culture. J. Econ. Entomol. 82: 236238. DuRant, J. A. 1985. Influence of temperature, relati ve humidity, date, and time of day on activity of the adult southern corn bill bug on corn. J. Agric. Entomol. 2(1): 20-26. Honek A. 1993. Intraspecific variation in body size a nd fecundity in in sects: a general relationship. Oikos 66 (3): 483-492 Jamjanya, T., and S. S. Quisenberry. 1988. Fall armyworm (Lepidoptera: Noctuidae) consumption and utilization of nine be rmudagrass. J. Econ. Entomol. 81: 697-704. Johnson-Cicalese, J. M. 1988. Billbugs pests of turfgrass: biology, host Range, and effect of endophyte-infected grasses. Master Thesis The State University of New Jersey. Johnson-Cicalese J. M. 1997. Developing turfgrass with enhanced insect resistance. TurfGrass TRENDS (6): 8.

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67 Johnson-Cicalese, J. M. and C. R. Funk. 1990. Additional host plants of four species of billbug found on New Jersey turfgrass. J. Am. Soc. Hortic. Sci. 115: 608-611. Johnson-Cicalese, J. M. and R. H. White. 1990. Effect of Acremonium endophyte on four species of billbug found on New Jersey turf. J. Am. Soc. Hort. Sci. 115(4): 602-604. Johnson-Cicalese, J. M., G. W. Wolfe, and C. R. Funk. 1990. Biology, distribution, and taxonomy of billbug turf pests (Coleoptera: Curculionidae). Environ. Entomol. 19(4): 1037-1046. Kamn, J. A. 1969. Biology of the billbug Sphenophorus venatus confluens a new pest of orchardgrass. J. Econ. Entomol. 62: 808-812. Kelsheimer, E. G. 1956. The hunting billbug, a serious pest of zoysiagrass. Proc. Florida Hort. Soc. 69: 415-418. Kindler, S. D. and E. J. Kinbacher. 1975. Differential reaction of Ke ntucky bluegrass cultivars to the bluegrass billbug, Sphenophorus parvulus Gyllenhal. Crop Sci. 15: 873-874. Kindler, S. D. and S. M. Spomer. 1986. Observations on the biology of the bluegrass billbug, Sphenophorus parvulus Gyllenhal (Coleoptera: Curculionid ae), in an eastern Nebraska sod field. J. Kansas Entomol. Soc. 56(1): 26-31. Kindler, S. D., S. M. Spomer, and E. J. Kinbacher. 1983. Further host range studies on the blue grass billbug, Sphenophorus parvulus Gyllenhal (Coleoptera: Curculionidae). Environ. Entomol. 12: 528-530 Koppenhfer, A. M., R. S. Cowles, and E. M. Fuzy. 2003. Effects of turfgrass endophytes (Clavicipitaceae: Ascomycetes) on white gr ub (Coleoptera: Scarabaeidae) larval development and field populati ons. Environ. Entomol. 32: 895. Kovitvadhi, K. and S. H. Kerr. 1968. Artificial diet for the zoysiagrass billbug, Sphenophorus venatus vestitus (Coleoptera: Curculionidae), and not es on its biology. Florida Entomol. 51(4): 247-250. Kunkel, B. A., P. S. Grewal, M. F. Quigley. 2003. A mechanism of acquired resistance against an entomopathogenic nematode by Agrotis ipsilon feeding on perennial ryegrass harboring a fungal endophyte. Bio. Control 29: 100-108. Matheny, E. L. 1981. Contrasting feeding habits of pest mole cricket species. J. Econ. Entomol. 74: 444-445. Montllor, C. B., E. A. Bernays, M. L. Cornelius. 1991. Responses of two Hymentoperan predators to surface chemistry of their prey : significance for an alkaloid-sequestering caterpillar. J. Chem Ecol. 17: 391-399.

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68 Morrill, W. L. and E. F. Suber. 1976. Biology and control of Sphenophorus coesiforns Gyllenhal (Coleoptera: Curculionidae) in bahiagrass. J. Georgia Entomol. Soc. 11: 283288. Murphy, J. A., S. Sun, and L. L. Betts. 1993. Endophyte-enhanced resistance to billbug (Coleoptera: Curculionidae), sod webworm (Lepidoptera: Pyralidae), and white grub (Coleoptera: Scarabaeidae) in tall fescue. Environ. Entomol. 22: 699-703. Nielson, D. C., K. H. Asay, and T. A. Jones. 1993. Bluegrass billbug feeding response to perennial triticeae grasses. J. Range Management. 46: 237-240. Niemczyk, H. D. and D. J. Shetlar. 2000. Destructive Turf Insects, 2nd ed. H.D.N. Books, Wooster, OH. Oliver, A. D. 1984. The hunting billbug one among the complex of turfgrass insect and pathogen problems. Amer. Lawn Appl. (March/Apr): 24-27. Peck, S. B. and M. C. Thomas. 1998. A distribution checklist of th e beetles (Coleoptera) of Florida. Arthropods of Florida and Neighboring Land Areas (16): 154-155. Potter, D. A. 1998. Billbugs, destructive tu rfgrass insects: biology, diagnosis, and control. Ann Arbor Press. Chelsea, MI. 170-177. Potter, D. A., C. G. Patterson, and C. T. Redmond. 1992. Influence of turfgrass species and tall fescue endophyte on feeding ecology of Japa nese beetle and southern masked chafer grubs (Coleoptera: Scarabaeidae). J. Econ. Entomol. 85: 900-909. Prestidge, R. A. and R. T. Gallagher. 1988. Endophyte fungus confers resistance to ryegrass: Argentine stem weevil larval studies. Eco. Entomol. 13: 429-435. Quisenberry, S. S. 1990 Plant resistance to insects and mites in forage and turf grasses. Florida Entomologist 73: 411-421. Quisenberry, S. S., and H. K. Wilson. 1985. Consumption and utilization of bermudagrass by fall armyworm ( Lepidoptera: Noctuidae) larvae. J. Econ. Entomol. 78: 820-824. Reigada C. and W. A.C. Godoy. 2005. Seasonal fecundity and body size in Chrysomya megacephala (Fabricius) (Diptera: Calliphoridae). Neotrop. Entomol. 34(2) Londrina Mar/Apr. 2005 Reinert, J. A., and P. Busey. 1984. Resistant varieties, pp. 35-40, in T. J. Walker (ed.). Mole crickets in Florida. Univ. Fl orida Agric. Exp. Stn Bull. 846. Reinert, J. A. and M. C. Engelke. 2001. Resistance to hunting billbug among Zoysia cultivars. http://esa.confex.com/esa/2001/techprogram/paper_1225.htm Texas A&M University, Research & Extension C enter Dallas. Texas. Complete.

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69 Richmond, D. S., H. D. Niemczyk, and D. J. Shetlar. 2000. Overseeding endophytic perennial ryegrass into stands of Kentucky bluegrass to manage bluegrass billbug (Coleoptera: Curculionidae). J. Econ. Entomol. 93(6); 1662-1668. Satterthwait, A. F. 1919. How to control billbugs destruct ive to cereal and forage crops. Farmers Bulletin 1003, USDA Washington, D.C. Satterthwait, A. F. 1931. Key to known pupae of the genus Calendra, with host-plant and distribution notes. Annals Entomol. Soc. Am. 24: 143-172. Schaffner, U., J. L. Boeve, H. Gfleller, U. P. Schlunegger. 1994 Sequestration of Veratrum alkaloids by specialist Rhadinoceraea nodicornis Konow (Hymenoptera: Tenthredinidae) and its ecoethological implicati ons. J. Chem. Ecol. 20: 3233-3250. Shetlar, D. J. 1991. Billbugs in turfgrass. Ohio Home, Yard, and Garden Facts, Ohio Coop. Ext. HYG 2502-91. 4 pp. Shetlar, D. J. 1995. Turfgrass insect and mite management, pp. 171-344. In T. L. Watschke, P. H. Dernoden, and D. J. Shetlar (eds.), Managing turfgrass pests. Lewis Publishers, Ann Arbor, Mich. 361 pp. Short, D. E. and E. A. Buss 2005. Bermudagrass mite. http://edis.ifas.ufl.edu/LH035 UF/IFAS Extension. Gainesville. Florida. Complete. Tashiro, H. 1987. Turfgrass Insects o f the United States and Canada. Cornell University Press, Ithaca, NY. Tashiro, H. and K. E. Personius. 1970. Current status of the bluegr ass billbug in its control in western New York home lawns. J. Econ. Entomol. 63: 23-29. Trenholm, L. E., J. L. Cisar, and J. B. Unruh. 2003. Bermudagrass for Florida lawns. http://edis.ifas.ufl.edu/LH007 UF/IFAS Extension. Gainesville. Florida. Com plete. Vaurie, P. 1951. Revision of the genus Calendra (formerly Sphenophorus ) in the United States and Mexico (Coleoptera: Curculionidae) Bull. Amer. Mus. Nat. Hist.93: 33-186. Wilson, D. and G. C. Carroll. 1997. Avoidance of high-endophyte space by gall-forming insect. Ecology. 78(7): 2153-2163. Woodruff, R. E. 2005. Hunting billbug, Sphenophorus venatus vestitus Chittenden (Insecta: Coleoptera: Curculionidae). http://edis.ifas.ufl.edu/IN364 UF/IFAS Extension. Gainesville. Florida. Co mplete.

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70 Young, F. B. 2002. Seasonal activity and biology of the hunting billbug, Sphenophorus venatus vestitus (Coleoptera: Curculionidae) in northwest Arkansas. M. S. thesis. University of Arkansas, Fayetteville.

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71 BIOGRAPHICAL SKETCH Ta-I Huang was born, grew up and attended school in Taipei, Taiwan. He studied forestry in the Chinese Culture University, w here he obtained his bachelors degree in 2003. He moved to the United States and studied at PALS English program at Rutgers University in 2005. He attended to the University of Florida, Entomology and Nematology Department in spring 2006 as a Graduate Research Associat e in the landscape entomology lab under Dr. Busss supervision where he started studying to ward his masters degree.