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University Microfilms International 300 N. ZEEB ROAD, ANN ARBOR, Ml 48106 18 BEDFORD ROW, LONDON WC1R 4EJ, ENGLAND 8022362

Wegner, G erald Sterling

BIONOMICS OF ATAENIUS SPRETULUS (COLEOPTERA: ) ON SOUTHERN OHIO GOLF COURSES

The Ohio State University Ph.D. 1980

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University M icrofilms International

200 N Z=£s *0. ANN Aa30S VII AS 1061313 ] 761-4700 BIONOMICS OF Ataenius spretulus

(COLEOPTERA: SCARABAEIDAE)

ON SOUTHERN OHIO GOLF COURSES

DISSERTATION

Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By Gerald Sterling Wegner, B.S., M.S.

*****

The Ohio State University

1980

Reading Committee: Approved By

Dr. Harry D. Niemczyk

Dr. David J. Horn

Dr. David G. Nielsen TAdVis* Dr. Keith J. Karnok tment of Entomology "The fear of the Lord is the

instruction for wisdom, and

before honor, comes humility."

Proverbs 15:33

ii ACKNOWLEDGMENTS

I wish to thank the following individuals for generously providing me with information from their respective fields of specialization: Oscar L. Cartwright, Entomologist Emeritus,

National Museum of Natural History, Smithsonian Institution;

H. Tashiro, Professor, Department of Entomology, New York

State Agricultural Experimental Station; Joseph Weaver,

Department of Entomology, West Virginia University; T. L.

Ladd, Japanese Laboratory, Ohio Agricultural Research and Development Center; Guilford S. Ide and Warren C.

Welbourn, Museum Curators, Acarology Laboratory, Department of

Entomology, Ohio State University; John L. Crites, Professor,

Department of Zoology, Ohio State University; Ronald L.

Stuckey and Roland L. Seymour, Professors, Department of

Botany, Ohio State University; James L. Caldwell, Professor, and T. Davis Sydnor, Associate Professor, Department of

Horticulture, Ohio State University; C. Wayne Ellett, Professor,

Department of Plant Pathology, Ohio State University; and

Douglas Streett, Department of Entomology, Ohio State

University.

I am grateful to the following organizations for their generous financial support of this project: the Musser

International Turfgrass Foundation; Ohio Turfgrass Foundation;

iii Golf Course Superintendents' Association of America? Cleveland

District Golf Association? and Golf Course Superintendents'

Associations of Greater Cincinnati, Central Ohio, Miami Valley,

Northern Ohio, Wisconsin, and Ontario, Canada.

I wish to individually thank the members of my graduate committee for their valuable advice and encouragement during the course of the study and in organizing the dissertation:

Harry D. Niemczyk, Professor, and David G. Nielsen, Associate

Professor, Department of Entomology, Ohio Agricultural

Research and Development Center? David J. Horn, Professor,

Department of Entomology, Ohio State University? and Keith J.

Karnok, Assistant Professor, Department of Agronomy, Ohio

State University. Many thanks are due the Ohio State

University Statistics Laboratory for assistance in analyzing data.

I am especially thankful to my wife, Debbie, for her loving cooperation and encouragement during the 3 year study and for typing the early drafts of the dissertation.

iv VITA

June 18; 1951...... Born - Lake Forest, IL.

1973...... B.S. in biology, Loyola University, Chicago, IL.

1973-1975...... Teaching Assistant, Department of Biology, Loyola University Chicago, IL.

197 5 ...... M.S. in biology Loyola University, Chicago, IL.

197 6 ...... Teaching Associate, Department of Entomology, The Ohio State University, Columbus, OH

1976-1979...... Research Associate, Department of Entomology, The Ohio Agricultural Research and Development Center, Wooster, OH.

PUBLICATIONS

Wegner, G. S . , & R.W. Hamilton. 1976. Effect of calcium sulfide on Chironomus riparius (Diptera: Chironomidae) egg hatchability. Environmental Entomology 5 (2) :256-8.

Wegner, G. S., & H. D. Niemczyk. 1979. The Ataenius of Ohio. Ohio Journal of Science, 79:249-55.

Niemczyk, H.D., & G.S. Wegner. 1979. Controlling the black turfgrass ataenius. Golf Course Management, 47:29-37.

Niemczyk, H.D., & G. S. Wegner. 1979. Life history and control of the black turfgrass ataenius. Ohio Report 64 (6) :85-88.

v VITA (Cont.)

FIELDS OF STUDY

Major Field: Entomology

Economic Entomology Turfgrass Entomology rearing and display techniques

vi TABLE OF CONTENTS

Page

ACKNOWLEDGEMENTS...... iii

VITA ...... v

LIST OF TABLES ...... xi

LIST OF FIGURES ...... xiv

INTRODUCTION ...... 1

LITERATURE REVIEW ...... 3

Taxonomy ...... 3 Geographic Distribution...... 5 Biology ...... 5 Similar Coleoptera ...... 8 Feeding Behavior ...... 8 Copulation and Oviposition ...... 12 Overwintering and Spring Activity .... 12 Baits and Attractants ...... 16 R e a r i n g ...... 18 Parasitism, Predation and Phoresy...... 20 Viral and Rickettsial Parasites .... 20 Bacterial Parasites...... 21 Microsporidial and Coccidial Parasites . 22 Gregarine Parasites...... 24 Nematode Parasites ...... 24 Insect Parasites ...... 25 Parasitic Fungi ...... 25 Insect Predators ...... 27 Vertebrate Predators ...... 27 Phoretic Acari ...... 28 P h e n o l o g y ...... 29 Sampling for Soil-Inhabiting .... 30

GENERAL METHODS AND MATERIALS ...... 31

INDIVIDUAL STUDIES ...... 33

Life History ...... 33 Methods and Materials ...... 33 R e s u l t s ...... 33

vii Page

INDIVIDUAL STUDIES (Cont.)

D i s c u s s i o n ...... 40 R e s u l t s ...... 46 Geographical Size of Population ...... 54 Dispersion Pattern of Population ...... 58 Flight and Insemination Studies ...... 66 Overwintering Studies ...... 79 Correlation of Plant Phenology, Temperature, and Humidity with Seasonal Life History of A. spretulus ...... 84 Associated Organisms ...... 90 Laboratory Oviposition studies ...... 96 Rearing and Developmental Studies ...... 100

GENERAL DISCUSSION ...... 108

SUMMARY ...... 110

REFERENCES CITED ...... 115

APPENDICES ...... 131

Appendix 1. Analysis of soil from sites sampled for overwintering survival of A. spretulus adults ...... 132

Appendix 2. Standard fairway treatments at 3 golf courses where A. spretulus life stage sampling was conducted from 1977 through 1978 . 133

Appendix 3. Number of each life stage of A. spretulus found per sampling occasion at TPCC during 1977 134

Appendix 4. Number of each life stage of A. spretulus found per sampling occasion at LCC during 1977 ...... 135

Appendix 5. Number of each life stage of A. spretulus found per sampling occasion at KCC during 1977 ...... 136

Appendix 6. Number of each life stage of A. spretulus found per sampling occasion at TPCC during 1978 137

Appendix 7. Sex ratio of A. spretulus adults collected from sampling sites at TPCC in 1977 . 138

viii Page

APPENDICES (Cont.)

Appendix 8. Sex ratio of A. spretulus adults 139 collected from sampling sites at LCC in 1977. . 140

Appendix 9. Sex ratio of A. spretulus adults collected from sampling sites at KCC in 1977. . 141

Appendix 10. Sex ratio of A. spretulus pupae collected from sampling sites at TPCC in 1977 . 142

Appendix 11. Sex ratio of A. spretulus pupae collected from sampling sites at LCC in 1977 . 143

Appendix 12. Sex ratio of A. spretulus pupae collected from sampling sites at KCC in 1977 . 144

Appendix 13. Sex ratio of A. spretulus adults collected from sampling sites at TPCC in 1978 . 145

Appendix 14. Sex ratio of A. spretulus pupae collected from sampling sites at TPCC in 1978 . 146

Appendix 15. Number of A. spretulus adults captured per day on rectangular screen sticky traps at TPCC during 1977 ...... 147

Appendix 16. Number of A. spretulus adults captured per day on rectangular screen sticky traps at LCC during 1977 148

Appendix 17. Number of A. spretulus adults captured per day on rectangular screen sticky traps at KCC during 1977 149

Appendix 18. Maximum, minimum and mean numbers of A. spretulus adults captured per day on rectangular screen sticky traps at TPCC, LCC and KCC during 1977 ...... 149

Appendix 19. Number of A. spretulus adults captured per day on 8-vaned sticky traps at TPCC during 1978 150

Appendix 20. Number of A. spretulus adults captured per day on 8-vaned sticky traps at KCC during 1978 ...... 151

ix Pages

APPENDICES (Cont.)

Appendix 21. Maximum, minimum and mean numbers of A. spretulus adults captured per day on 8-vaned sticky traps at TPCC and KCC during 1978...... 152

Appendix 22. Number of A. spretulus adults captured per day in author-made suction black light trap at TPCC during 1977 .... 153

Appendix 23. Number of A. spretulus adults captured per day in Will-o'-the-Wisp ® suction black light trap at LCC during 1977 . 154

Appendix 24. Number of A. spretulus adults captured per day in Will-o’-the-Wisp ® suction black light trap at KCC during 1977 . 155

Appendix 25. Mean number of A. spretulus adults captured per day in suction black light traps at TPCC, LCC and KCC durina 1977 155

Appendix 26. Number of A. spretulus adults captured per day in Will-o1-the-Wisp ® suction black light traps at TPCC during 1978 156

Appendix 27. Number of A. spretulus adults captured per day in Will-o'-the-Wisp ® suction black light trap at KCC during 1978 157

Appendix 28. Mean number of A. spretulus ^ adults captured per day in Will-o'-the Wisp^-' suction black light traps at TPCC and KCC during 1978 ...... 157

Appendix 29. Data tabulated for analysis of variance to test effect of site, month and sex on survivorship of overwintering adults of A. sp r e t u l u s ...... 159

Appendix 30. Nutrient mixture for hydroponic culture of turfgrass (after Hoagland and Arnon 1950 and Christians 1977) ...... 160

Appendix 31. Artificial diet tested on larvae of A. spretulus (modified from Yamamoto 1969 and Bell and Joachim 1976) ...... 161

x LIST OF TABLES

TABLE Page

1. Dimensions of A. spretulus life stages, based on measurements (in mm) of 20 field- collected specimens per life stage...... 34

2. Summary of A. spretulus life stage sampling record at TPCC, LCC, and KCC during 1977 - 1978 ...... 47

3* X^comparison of A. spretulus infestations in TPCC sampling sites 14 and 17 by age class, using 1977 d a t a ...... 56

4. X comparison of A. spretulus infestations in TPCC sampling sites 17 (1977 data) and 14B (L978 data) by age c l a s s ...... 57 2 5. % comparison of A. spretulus infestations in TPCC sampling sites 14 (1977 data) and 14A (1978 data) by age c l a s s ...... 57 2 6. X comparison of A. spretulus infestations in TPCC sampling sites 14A and 14B by age class, using 1978 d a t a ...... 59 2 7. X comparison of A. spretulus infestations in sampling sites at TPCC, LCC, and KCC by age class, using 1977 data pooled by course ...... 59

8. Fit of A. spretulus population from TPCC 14 to Poisson and negative binomial distributions by age class, using 1977 d a t a ...... 63

9. Fit of A. spretulus population from TPCC 17 to Poisson and negative binomial distributions by age class, using 1977 d a t a ...... 63

xi TABLE Pages

10. Pit of A. spretulus population at TPCC (pooled 1977 data from sampling sites 14 and 17) to Poisson and negative binomial distributions by age class ...... 64

11. Fit of A. spretulus population at LCC (pooled 1977 data from*sampling sites 11 and 16) to Poisson and negative binomial distributions by age class ...... 64

12. Fit of A. spretulus population at KCC (pooled 1977 data from sampling sites 4 and 6) to Poisson and negative binomial distributions by age class ...... 64

13. Captures of A. spretulus on 8-vaned sticky traps at two heights in 1978. Significance between captures at different levels and relationship to average monthly tempera­ ture are given ...... 73

14. Percentage of inseminated A. spretulus females captured in black light traps at TPCC, LCC, and KCC during 1977...... 76

15. Percentage of inseminated A. spretulus females captured in black light traps at TPCC and KCC during 1978 76

16. Effect of site, month, and sex on survivorship of overwintering A. spretulus adults near Cincinnati, O h i o ...... 77

17. Effect of site and month on survivorship of overwintering A. spretulus adults near Cincinnati, O h i o ...... 77

18. Day degree basis of A. spretulus biology and associated plant phenology ...... 88

19. Incidence of milky spore-diseased A. spretulus at TPCC, LCC, and KCC during 1977 and 1978 . . 92

20. Suitability of various substrates on egg maturation in and oviposition by inseminated, overwintered females of A. spretulus at 23+2 ° C ...... 97

xii TABLE Pages

21. Life table for a laboratory-reared generation of A. spretulus held at 23+2 C, 68+12* R.H., starting with field-collected eggs and using data from life stage samples and overwinter­ ing studies to estimate time span and survivorship from the prereproductive to the reproductive adult interval ...... 105

xiii LIST OF FIGURES

FIGURE Pages

1. Egg cluster of A. spretulus ...... 35

2. First, second, and third instar A. spretulus . . 35

3. Rastral pattern of mature A. spretulus larva . . . 37

4. Scanning electron micrograph of hamate setae on raster of mature A. spretulus larva .... 37

5. Dorsal aspect of left larval mandible of A. spretulus, showing long seta (a) and short seta (b)...... 38

6. Female (a) and male (b) A. spretulus pupae. Arrow indicates aedeagal protuberance of male. 39

7. A. spretulus a d u l t ...... 39 2 8. Typical 558.15 m sampling site, subdividied into plots (a) and quadrats (b), in which unrestricted random sampling for life stages of A. spretulus was conducted ...... 44

9. Seasonal occurrence and abundance of A. spretulus life stages in sampling sites at TPCC during 1977-78 ...... 49

10. Seasonal occurrence and abundance of A. spretulus life stages in sampling sites at LCC and KCC during 1977...... 50

11. Rectangular, screen sticky trap used to monitor seasonal flight activity of A. spretulus in 1977 ...... 67

12. Eight-vaned sticky trap used to monitory seasonal flight activity of A. spretulus in 1978 ...... 67

13. Author-made suction black light trap used in 1977 to monitor daily and seasonal A. spretulus flight activity...... ' . 70

xiv FIGURE Page

14. Modified Will-o'-the-Wisp Buglite Fish Feeder® suction black light trap used in 1977 and 1978 to monitor daily and seasonal A. spretulus flight activity ...... 70

15. Seasonal occurrence and abundance of A. spretulus adults on rectangular screen sticky traps (averaged per day) at TPCC, LCC, and KCC during 1977 ...... 72

16. Seasonal occurrence and abundance of A. spretulus adults on 8-vaned sticky traps (averaged per day) at TPCC and KCC during 1978 72

17. Seasonal occurrence and abundance of A. spretulus adults in author-made and Will-o'-the-Wisp® suction black light traps (averaged per day) at TPCC, LCC, and KCC during 1977 ...... 75

18. Seasonal occurrence and abundance of A. spretulus adults in Will-o1-the-Wisp®suction black light traps (averaged per day) at TPCC and KCC during 1978 75

19. Typical overwintering site of A. spretulus adults at perimeter of golf course ...... 81

20. A. spretulus adult overwintering site in rough between fairways at a golf c o u r s e ...... 81

21. Protective encasement with circular chart, 7- day recorder used to record soil temperature 5 cm deep in a sampling site at KCC during 1978 ...... 86

22. Milky spore diseased larva of A. spretulus (a) compared to healthy larva (b)...... 93

23. Cephaline eugregarine sporont found in third instar larva of A. spretulus: shape of living sporont (a) and photo-micrograph of preserved specimen (b) ...... 93

24. Ovaries of A. spretulus with eggs in late stage of development ...... 99

25. Ovaries of A. spretulus showing ovarioles with eggs in early stage of development...... 99

26. Observation rearing unit (a) with 10 trans­ parent rearing troughs (b) for viewing developing larvae of A. spretulus ...... 101

xv FIGURE Page

27. Hydroponic cultures of Poa annua L. used as a source of root-diet for laboratory rearing larvae of A. s p r e t u l u s ...... 101

xvi I. INTRODUCTION

The black turfgrass ataenius, Ataenius spretulus

(Haldeman) \ is a pest of turfgrasses on golf courses and in the landscape, primarily in the eastern and midwestern United

States. Attempts to control this pest have often failed because control measures were not employed at the proper time or improper control tactics were used.

A. spretulus was identified and recognized as an important pest over much of its range during the early 1970's following much misdiagnosis of the problem and poor pest management practices. The turfgrass condition resulting from larval feeding damage was often improperly diagnosed as heat stress by golf course personnel. Niemczyk and Dunbar (1976) found evidence of cyclodiene insecticide resistance in several

A. spretulus populations. They suggested resistance may have developed as a result of widespread use of cyclodiene insecti­ cides on golf courses.

The present study was inspired by the need for biological information on A. spretulus pertinent to management. The magnitude of the problem induced golf course and turfgrass management organizations in several states to help sponsor full-time research on the subject. An intensive 3-year study

^(Coleoptera: Scarabaeidae) of A. spretulus biology and seasonal history was conducted at several Cincinnati-area (Hamilton and Clermont counties) golf courses during 1976-1978. II. LITERATURE REVIEW

TAXONOMY

A. spretulus (: Eupariini) was misidentified

frequently for nearly 100 years. Haldeman (1848) originally described the beetle and named it Aphodius spretulus. LeConte

(1863) named it Euparia spretulus (Hald.). Horn (1887) mistakenly categorized specimens of A. spretulus as examples of two similar species, Ataenius striqatus (Say) and A. coanatus

(LeConte). Later, Fall (1930) elucidated the distinguishing characters of A. striqatus and A. coanatus, but erroneously believed A. spretulus to be a new species, and assigned it the synonym A. consors. Hinton (19 34) changed the name from

A. consors Fall to A. falli Hinton. More recently, Cartwright

(1943, 1948, 1974) established the correct Latin binomial, elucidated external differences between male and female , and provided a dependable taxonomic basis for distinguishing

A. spretulus from morphologically similar species within the striqatus complex.

A condensed version of Cartwright's description of A. spretulus follows: Length 3.6 to 5.5 mm; width 1.7 to 2.4 mm.

Elongate-oblong, somewhat convex, shining black, legs and clypeas broadly rounded on each side of wide, shallow median

3 emargination. Surface of genae feebly, transversely wrinkled anteriorly, finely, closely punctate above middle. Pronotum with evenly spaced, fine punctures throughout and very irregularly placed, scattered, coarse punctures intermixed.

Apical fringe of metatibia usually with a group fo 5 setae and an intervening seta between the accessory spine and spurs.

First segment of metatarsus longer than long spur and longer than the following three tarsal segments combined. Males with patch of close, setigerous punctures on the metasternal disc, females without. Males also have a longer pygidium with a much wider apical lip than females.

The larva of A. spretulus may have been described on two occasions without the investigators knowing which Ataenius species they were describing. Hoffman (1935) described the larva of a species which he though was A. coqnatus. The adults reared from his larvae had been misidentified as A. coqnatus and are now thought to be A. spretulus. Jerath (1960) described three species of eupariine larvae which he designated

"Ataenius spp. 1, 2, and 3". All larvae were found in association with adults of A. spretulus. One of these unknown species was probably the larva of A. spretulus.

Hoffman (1935) may have described the prepupa and pupa of A. spretulus in the same unknowing way that he described the larva. He gave no method for distinguishing male and female pupae. The method given by Tashiro, et al. (1969) for sexing pupae of the European chafer, Rhizotroqus (= Amphimallon) malalis Razoumowsky, applies to pupae of other Scarabaeidae, including A. spretulus. 5

GEOGRAPHIC DISTRIBUTION

Cartwright (1974) reported that A. spretulus has been

found in all but nine of the United States and in Ontario,

Canada. There is no record of the beetle from Alaska,

Arizona, Hawaii, Maine, Montana, Nevada, Oregon, Vermont, or

Washington at this writing. Niemczyk and Wegner (19 79)

reported that economically important populations of A.

spretulus exist in parts of 23 states (Colorado, Connecticut,

Delaware, Illinois, Indiana, Iowa, Kansas, Kentucky, Maryland,

Massachusetts, Michigan, Minnesota, Missouri, Nebraska, New

Hampshire, New Jersey, New York, Ohio, Pennsylvania, Rhode

Island, Virginia, West Virginia, Wisconsin) and Ontario,

Canada.

Cartwright (1974) listed the following Ohio county and

city () locations respectively, in which A. spretulus had

been collected: Ashtabula (Conneaut), Athens (Athens),

Champaign, Cuyahoga (Cleveland), Erie, Franklin, Hardin (Ada),

Henry (Holgate), Lake (Painesville), Ottawa, Preble (West

Alexandria), Summit (Hudson), and Washington. Drees (1977)

reported A. spretulus from Adams, Clermont, Darke, Fairfield,

Hamilton, Lorain, Stark, and Wayne counties, Ohio. There is

no report of A. spretulus from outside the continental

United States and Canada.

BIOLOGY

Little has been written on the biology of A. spretulus.

Forbes (1905) may have made the first pertinent observations when he reported taking an adult Ataenius coanatus (probably

A. spretulus or A. striqatus) from the wet, decaying husks

of a fallen ear of corn at Farina, Illinois. He observed

the emergence of a second beetle from a pupa found among corn

roots on 21 July, 1905. Blatchley (1910) reported large numbers of adult A. coqnatus (probably A. spretulus or A. strigatus) hibernating in and beneath dry cow dung in Indiana.

He noted that beetles flew during warm sunny days in winter, frequented fungi, and were attracted to electric lights. In a later paper on the Scarabaeidae of Florida, Blatchley (1928) wrote that the species occurred throughout the state, sometimes by the hundreds beneath piles of decaying Chara sp. and weed debris along ditches.

Several sources in the literature suggest that A. spretulus larvae adapt readily to a diet consisting mainly of turfgrass roots: Hoffman (1935) reported A. coqnatus (probably A. spretulus) as a new pest of golf courses following an infestation in St. Paul, Minnesota. Larvae caused severe damage by killing grass in patches on greens and fairways.

All stages were most abundant immediately below the turf; no grubs or pupae were found below a soil depth of 5 inches

(12.7 cm). Samples from infested turfgrass yielded 60 to 2 2 2 116 larvae/ft (per 929 cm ) and 32 to 42 pupae/ft . Hoffman noted adults were taken at light traps as early as May 1. He deduced that oviposition occurred in late May or June and that adult emergence was completed by 27 August. The species were observed to overwinter only in the adult stage. In September, Hoffmann discovered a large number of adults

"awaiting winter" within the upper 6 inches (15.24 cm) of waste piles composed of milorganite and moist grass cuttings from golf course greens. Hoffmann concluded there was only one generation per year in Minnesota.

Frost (1966) reported taking an A. spretulus adult in a black light trap on 22 January, 1963 in Highlands County,

Florida. Woodruff (1973) wrote that A. spretulus adults occasionally were found in great numbers around electric lights in southern Florida. Adults were collected there every month of the year. Frost's and Woodruff's reports suggest that A. spretulus development may be continuous in the far southern portion of its range. Woodruff also men­ tioned that he had collected a few beetles in cow dung and one in deer droppings.

Kawanishi, et al. (1974) encountered larvae of A. spretulus at two golf courses in New York, one in Monroe

County and the other in Orange County, during 1969-1971.

Their observations supported Hoffman's report of one generation per year. Niemczyk (1975, 1977) Niemczyk and Dunbar (1976), and Niemczyk and Wegner (19 79) described A. spretulus larval damage to annual bluegrass (Poa annua L.) bluegrass (P. pratensis L.), and bentgrass (Aarostis spp.) on golf courses in Canada and the United States. Larval populations estimated 2 2 in excess of 300/ft (per 929 cm ) occurred in fairways at one country club in Cincinnati, Ohio in 1973. Niemczyk and Dunbar (1976) provided evidence for two generations of A. spretulus per year in both Ohio and Connecticut. In Ohio, overwintered adults were first observed flying in early May.

First generation adults were observed from early July to early August; second generation adults were first seen in early September. In Connecticut, first generation adults were observed in mid- to late July, and second generation adults emerged in mid-September.

Weaver and Hacker (1978) described A. spretulus larval damage to golf course fairways in the West Virginia counties of Putnam, Cabell, Kanawha, and Wood. They found evidence for two generations per year in West Virginia. The first oviposition period lasted from early May to mid-June; mature larvae were present from mid- to late June; first generation adults began emerging in late June. The second oviposition period lasted from early July to mid-September; second generation adults began appearing in early Agust. Overwinter­ ing adults were most commonly found in wooded areas where there was an accumulation of leaf litter and the soil was loose and well-drained.

SIMILAR COLEOPTERA

FEEDING BEHAVIOR

Beetles in the genus Aphodius are similar in appearance and behavior to those in Ataenius. A few species of Aphodius have exhibited the same dichotomous diet perferences attributed to A. spretulus. Larvae of both genera have been observed feeding on dung or humus on some occasions and plant roots at other times. Downes (1928) reported Aphodius paradalis

LeConte as a root-feeding turfgrass pest in lawns in British

Columbia, Canada. Ritcher and Morrison (1955), Jerath and

Ritcher (1959), and Jerath (1960) reported finding A. pardalis 2 2 larvae in abundance (240 to 644/ft or per 929 cm ) beneath damaged patches of turfgrass on a golf course in Eugene, Oregon in 1954. Jerath (1960) also collected larvae of this beetle beneath decaying lawn grass at San Francisco, California in

1935. Records of this species as a pest are rare enough to suggest that whole populations of A. pardalis may subsist, inconspicuously and harmlessly, on dung or decaying organic matter.

Jerath and Ritcher (1959) observed larvae of Aphodius aranarius (L.) feeding on sprouting maize seed in Minnesota, and on grass roots in Oregon. Niemczyk (1978) found A. qranarius larvae in great abundance beneath damaged P. annua on golf course fairways in Boulder, Colorado and Detroit,

Michigan. In reference to A. aranarius as a non-injurious species, Blatchley (1910), Mohr (1943), Jerath (1960), and

Woodruff (1973) found larvae of this species associated with cow dung in Indiana, Illinois, Kentucky, and Florida, re spe ct i ve ly.

Swan (1934), Given (1950), and Jerath and Ritcher (1959) reported Aphodius howitti Hope (=A. tasmaniae Hope) as a pest of pasture plants in parts of South Australia, Tasmania, and

New South Wales. Cumpston (19 41) observed that areas rich in sheep dung were attractive to ovipositing females. Jerath and Ritcher (1959) reported that Aphodius pseudotasmaniae Given fed on roots of pasture plants in

Tasmania, Aphodius fimetarius (L.) was a pest of potatoes in

Bremen, Germany, Aphodius contaminatus Herbst was collected in golf course turf in Oregon, Aphodius distinctus Muller was injurious to mint roots in Quincy, Washington, and Aphodius hamatus Say caused damage to pasture turf in Ruby Valley,

Nevada. Reports of dung or humus-feeding behavior do not exist for all of the above species of Aphodius. However,

Sanders and Dobson (1966) collected both larvae and adults of

A. fimetarius in bovine manure in Indiana; Woodruff (1973) stated that A. fimetarius is a common species in cow and horse dung in Florida; Mohr (1943) found both A. fimetarius and

A. distinctus in cattle droppings in pastures at Urbana,

Illinois. The other aforementioned species may feed on and develop within dung or humus under certain circumstances. The circumstances dictating the dietary preference of some scara- baeid species are not well understood, but one explanation was offered by Cumpston (1941). He observed that phytophagous scarab larvae actually feed on both living roots and on humus or some other form of decaying organic matter, and suggested that, in certain cases, larvae may favor root tissue only if there is a deficiency of the preferred decayed matter in the immediate environment. Cumpston noted that larvae of phytophagous scarabs, such as Heteronvchus sanctae-helenae

Blanchard (Dynastinae) and Anodontonvx tetricus Blackburn

(Melolonthinae) have been reared in the laboratory on humus 11

alone. According to Jepson (1956), larvae of Heteronvchus

tenuistriatus Fairmaire usually feed on decaying roots and

detritus but, when numerous, damage roots of sugar cane in

Tanzania.

Blatchley and Leng (1916) wrote that weevils of the

genus Hyperodes (Curculionidae: Curculioninae) occur

beneath cover on sandy or muddy ground near water, feed upon

semi-aquatic plants, and probably are not injurious to

agriculture. Over fifty years later, Schread (1970) reported

that root-feeding H. anthracinus (Dietz) and H. maculicollis

(Kirby) had injured annual bluegrass fairways on seven golf

courses in Connecticut. Cameron (1970) and Cameron and

Johnson (1971a, b) reported similar damage by these two weevils at golf courses in New York and Pennsylvania.

Cameron and Johnson found that on Long Island, Hvperodes

females deposit their egges between leaf sheaths of annual bluegrass during April and May. Larvae feed on grass stems, pass through five instars, and pupate about 0.25 inch (0.64 cm)

deep in the soil. Mature adults become active at dusk and may be found at lights at certain times during the summer.

Schread (1970) claimed one generation per year for H. anthracinus

and H. maculicollis in Connecticut, with adults emerging in

late June and July, whereupon they enter reproductive diapause until the following spring. Cameron (1970), Cameron and Johnson (1971a, b), and Tashiro (1976) suggested there may be two generations in New York, with the second generation occurring from late June to late September. 12

COPULATION AND OVIPOSITION

The literature contains few recorded observations of

aphodiine beetles in copula. Woodruff (1973) found

several mating pairs of Aphodius crassulus Horn in horse dung in a Florida hammock, and two copulating pairs of

Pleurophorus lonoulus Cartwright near gasoline pump lights at a service station. Apparently there is no scarcity of information on copulation in some other scarabaeid subfamilies.

Davis (1916) and Ritcher (1958) wrote that adults of several species of Liovrus (Dynastinae) copulate underground.

Smith and Hadley (1926) and Fleming (1972) provided detailed accounts of Japanese beetles, Popillia iaponica Newman

(Rutelinae) in copula on foliage. Tashiro, et al. (1969) observed adult Rhizotroqus maialis (Razoumowsky)

(Melolonthinae) copulating on tree foliage. Ritcher (1958) reported that Geotrupes (Geotrupinae) and Pleocoma (Pleoco- minae) mate in burrows in the soil.

Univoltine scarabaeids which overwinter as adults usually mate in the fall and oviposit the following spring.

Ritcher (1958) found this to be the case for species of

Geotrupes. Pleocoma, Anomala, and Aphodius. Davis (1916) found the same for Licvrus spp.

OVERWINTERING AND SPRING ACTIVITY

As stated earlier, A. spretu]us adults overwinter beneath cow dung (Blatchley, 1910), in and beneath compost

(Hoffmann, 1935), beneath leaf litter, and in loose soil 13 within wooded areas (Weaver and. Hacker, 1978). Species in the genera Geotrupes, Pleocoma, Anomala, Aphodius, and Liqyrus overwinter in similar locations (Ritcher, 1958 and Davis

1916).

A number of scarabaeids overwinter as larvae, including members of the subfamilies Rutelinae (Smith & Hadley, 1926;

Fleming, 1972) and Melolonthinae (McColloch & Hayes, 1923;

Tashiro et al., 1969).

Benton and Crump (1979) observed overwintering behavior of Coleomegilla maculata (DeGeer) (Coccinellidae). Adults were stimulated to seek overwintering sites on the basis of shortening photoperiod and exhibited hygrotactic and anemotac- tic behavior during emigration flights. Beetles aggregated at the bases of prominent objects, especially tree trunks, on a south-facing slope. Adults apprently entered a state of ateleodiapause beneath leaf litter and undergrowth. Over­ wintering survivorship was 97-99%.

Schread (1970) , Cameron and Johnson (1971a, b) and

Tashiro (1976) conducted overwinterinq studies with the weevils h . anthracinus and H. maculicollis. These investigators determined that these species overwinter as adults in protected areas on or near golf courses, particularly in tufts of fescue and among leaves and debris at the edge of wooded areas. Tashiro (1976) found up to 40 weevils 929 cm in such areas. Overwintered adults became active soon after thawing temperatures predominated in spring. Some adults flew 14

from overwintering sites to suitable breeding sites while others emigrated on the ground.

Studies on overwintering and spring flight activity have also been conducted with bark beetles (Scolytidae) since they are economically important on a variety of evergreen and hardwood trees. Reid (1963) reported that adult overwintering mortaility in mountain pine beetle, Dendroctonus ponderosae

Hopkins, was a function of altitude. He found, nearly 100% mortality among beetles overwintering in Pinus above the snow line and about 50% mortality below the snow line in British

Columbia. Frye, et al. (1974) found that a severe cold period (-40 °c) during the week of 1 January, 1971 in Arizona, resulted in 28% mortality of hibernating adult spruce beetle, Dendroctonus rufioennis (Kirby), located in the bark and wood at the bases of spruce trees, below the snow level.

He noted that bark beetle susceptibility to cold temperature appears to be greatest in early winter when moderate temper­ atures are followed by a sudden drop to sub-zero temperatures.

Watson (1971) stored D. ponderosae adults at 1 °c in plastic bags containing mostened excelsior. Under these conditions,

85% of the beetles survived after 30 days; female survivor­ ship in storage was greater than males. Safranyik (19 76) observed that larger individuals of both sexes of D. ponderosae generally survived longer than smaller individuals when stored at 1 - 2 °C. He also noted the maleifemale ratio of the surviving beetles decreased with increased duration of storage. 15

The onset of spring emergence and flight behavior by- overwintering adult bark beetles is dependent on a favorable combination of light intensity and temperature. McMullen and

Atkins (1962) noted that the Douglas fir beetle, Dendroctonus pseudotsuqae Hopkins, flies when the maximum air temperature reaches 65 - 70 °f (18.3 - 21.1 °c) for a few days. Zethner-

Moller and Rudinsky (1967) found that emergence and flight of

Hylastes niarinus Mannerheim were at their maximum in late

April and early May, occurring between 61 and 75 °F (16.1 -

23.9 °C). They also found flight behavior peaked at light 3 intensities between 300 and 1200 footcandles (3.23*10 4 1.29*10 lux), during the late afternoon or early evening.

Rudinsky and Schneider (1968) recorded a lower temperature threshold of 58 °p (14.4 °C) and an upper light intensity threshold of 2000 footcandles (2.15-10^ lux) for flight activity of Gnathotrichus retusus LeConte and G. sulcatus

LeConte.

Mass flight of bark beetles in spring is considered to have a dual purpose. According to Rudinsky (1962), a short flight period serves to disperse the population, providing for the location of a suitable host tree. Additionally, Graham

(1959) suggested, "the activity of flight may play an important role in abolishing the photopositive response... permitting the beetles to react to contact and olfactory stimuli." 16

BAITS AND ATTRACTANTS

Woodruff (1973) cited a number of situations in which aphodiine beetles were attracted to liquid baits of various kinds. He collected A. spretulus adults in Florida in

McPhail fruit fly traps baited with a fermenting mixture containing pineapple juice; Ataenius platensis (Blanchard) in traps baited with rotting cantaloupe and dead fish;

Aphodius bicolor Say, Aphodius campestris Blatchley, and

Aphodius haldemani Horn in malt bait traps, in some cases, containing a few drops of propionic acid. Sex or aggregation pheromones have not been reported for members of the

Aphodiinae.

Most of the information available on synthetic and natural baits and attractants for scarabaeids applies to the economically important species of the Melolonthinae and

Rutelinae. Tashiro, et al. (1969) found that a 3:1 ratio

(v/v) of Java citronella oil and eugenol was the most consis­ tently attractive mixture of aromatic oils to adults of the

European chafer, R. maialis. They also reported the single most attractive synthetic compound to R. maialis was butyl sorbate and that flight traps painted black and red attracted

7% more flying adults than yellow, 21% more than white, and

22% more than blue. Traps containing virgin females or their extracts were noc attractive. Smith and Hadley (1926) observed that virgin female P. iaponica were highly attractive to males, often stimulating great numbers of males simultane­ ously to attempt copulation with a single female. Fleming, 17 et al. (1940) wrote that yellow flight traps were 15-65% more attractive to P. iaponica adults than traps painted silver, white, red, blue, green, or mixtures of these colors. Ladd, et al. (1975) reported that a trinary mixture of phyenthyl propionate (PEP), eugenol, and geraniol was most attractive to adults of P. iaponica.

Pheromonal communication between individuals of a popu­ lation has been demonstrated in some beetles. Pheromone- initiated mass attack is an aspect of bark beetle (Scolytidae) biology which has received much attention. This behavior is expressed during the spring flight period of the overwintered beetles and is characterized by three distinct phases of aggregation (Vite' and Pitman 1968; Wood, et al. 1970):

(1) detection and selection of suitable host trees; (2) mass attack and colonization; and (3) terminal attack on the target tree and possible mass attack of adjacent trees.

Onset of mass attack is triggered by release of an aggregation pheromone by one or more scout beetles which have located suitable host trees. In the genus Dendroctonus, scout beetles are all female (Person, 19 31) and the aggregation pheromones released are highly attractive to both sexes of their species

(Pitman and Vite', 1969; Bendard, et a l . 1969) . In the genus

I p s , males are the scouts (Anderson, 1948) and the aggregation pheromones they release are attractive only to conspecific females (Vite' and Pitman, 1968; Wood, et al. 1970). 18

The most reliable methods employed by early investigators

for studying bark beetle aggregation behavior were outdoor

baits and laboratory ofactometer tests. McMullen and Atkins

(1962) and Wood, et al. (1970) used freshly-cut logs of host

conifers as baits to attract scout beetles on Dendroctonus and

I p s and to observe mass attack. In the laboratory, Wood (1962)

used a multiple choice olfactometer with samples of host wood,

virgin males and females, and various controls as choices, to

elucidate the pattern of attraction that occurs during mass

attack.

REARING

There is little published information on rearing aphodiine

beetles. Apparently no one has established a continuous

rearing technique for any species in this subfamily. However,

a number of investigators have reared A. spretulus larvae

to adults. Hoffmann (1935) reared A. spretulus larvae to

adults in salve tins and jelly glasses filled with a soil-

milorganite mixture topped with sod. Cartwright (1974)

reared at least ten species of Ataenius from larva to adult in

containers filled with various mixtures of humus, soil, sand,

and dung. A few researchers have been able to induce certain

aphodiine species to oviposit in captivity and have reared

the resulting larvae to adults. Ritcher (1966) obtained

Aphodius larvae by confining adults in pint jars of fresh manure. Cartwright (1974) achieved some oviposition and

rearing of Ataenius by placing adults in pint jars filled with moist, sterile sand mixed with small amounts of finely ground dry breakfast cereal. Rearing methods for other scarabaeids have been published

Girault (1914) and Davis (1915) devised several methods for

rearing phytophagous scarabaeid larvae in the subfamilies

Dynastinae, Melolonthinae, and Rutelinae. Their rearing

technique utilizing terra-cotta flower pots is still used

today. Davis introduced copulating pairs of beetles to 15

inch (38.1 cm) diameter flower pots filled with rich, sifted

soil seeded with timothy (Phleum pratense L.) and bluegrass

(P. pratensis). The pots also contained pieces of old corn­

stalk as a source of decaying vegetable matter. The pots were

covered with cylindrical or conical, 20-mesh wire tops to

prevent adult escape. Introduced beetles oviposited in the

pots, and resulting larvae commonly developed into adults.

Occasionally pots had to be reseeded.

Sanders and Fracker (1916) developed a special observa­

tion rearing chamber for monitoring larval development of

Lachnosterna spp. (Melolonthinae) in the soil. They con­

structed each rearing unit from two panes of glass mounted

< 1.3 cm apart in an open top wooden frame. This unit was

filled with soil, and seeded with turfgrass or corn according to the known preference of the species being reared. Tashiro,

et al. (1969) induced oviposition by field-collected, gravid

R. maialis females in moist, screened loamy soil in 30 lb.

(13.64 kg) freezer tins. 20

PARASITISM, PREDATION, AND PHORESY

Viral and Rickettsial Parasites

Most of the work on viral diseases of Scarabaeidae has been done on Austrialian fauna. Steinhaus and Leutenegger

(1963) discovered an icosahedral virus in the larval fat body of the pruinose scarab, Sericesthis pruinosa (Dalman)

(Melolonthinae) in New South Wales. This same Sericesthis iridescent virus (SIV) also utilized Tenebrio molitor L.

(Tenebrionidae), Aariotes obscurus (L.) (Elateridae), and

Melolontha sp. as hosts in laboratory studies. Goodwin and

Filshie (1969) coined the term "spheroidosis" virus to describe a particle they isolated from larval fat body to

Othnonius batesi Olliff (Melolonthinae) in New South Wales.

Goodwin and Filshie (1975) found this parasite, which they placed in the Entomopoxvirus (D)/*:*/:X/X:I/0 (Vagoiavirus) group, in larvae of Aphodius tasmaniae Hope and Dermolepida albohirtum (Waterhouse) (Melolonthinae). Kalmakoff, et al.

(1972) isolated an iridescent virus, which they designated

CzIV, from larvae of Costelvtra zealandica (White)

(Melolonthinae) in New Zealand. Lipa and Bartkowski (1972) discovered an entomopoxvirus in diseased larvae of Geotrupes silvaticus (L.) in Poland. Krieg and Huger (1960) found that a nonocculuded virus, Wassersucht virus of Coleoptera, causes disease in some species of Melolontha. They noted that infected larvae appeared transparent, biege and waxy in the late stages of disease.

Hurpin (1968) reported the rickettsia Rickettsiella melolonthae (Rickettsiaceae: Wolbachieae) as a parasite of 21 both Melolontha hippocastani L . and Melolontha melolontha L ..

Vaught (1974) listed four rickettsia-scarabaeid parasite- host relationships: Rickettsiella popillae from P. iaponica.

R. melolonthae from M. melolontha, and two Rickettsiella spp. from Aphodius tasmaniae and Sericesthis niorolineata

(Boisduval) . Vaughn explained that the disease produced by

Rickettsiella spp. infecting the fat body of a white grub host is often called "blue disease" because diseased grubs are bluish.

Bacterial Parasites

Three strains of milky disease caused by Bacillus spp.

(Bacillaceae), have been isolated from A. spretulus.

Kawanishi, et al. (1974) discovered the first of these bacilli in ca. 66% of the grubs examined from a golf course in Monroe

County, New York. They found that the endospore of this unnamed milky disease organism is structurally similar to, yet distinct from, the endospore formed by known strains of

Bacillus lentimorbus Dutky and 3. popillae Dutky.

Splittstoesser and Tashiro (1977) discovered two additional

Bacillus strains in ca. 25% of A. spretulus larvae examined from the same golf course in New York in 1975. The two latter bacilli were distinct from each other and from the first in both spore size and morphology. The hemolymph of diseased grubs contained spores of only one of the three Bacillus strains. 22

Neither B. popillae nor B. lentimorbus have been reported

from larvae of A. spretulus, although strains of one or both

species are known to infect P. iaponica larvae (Dutky, 1940:

Cory, 1940; White, 1940; Beard, 1944), R. maialis (Tashiro &

White, 1954; Tashiro, et al. 1969), M. melolontha (Hurpin &

Robert, 1968, 1972) and Cvclocephala borealis Arrow

(Dynastinae) (Harris, 1959).

Microsporidial and Coccidial Parasites

Miscrosporidians (Protozoa: Cnidospora: Microsporea) are intracellular parasites of fish and (Barnes,

1968). Most of the microsporidian parasites reported from

Coleoptera belong to the family Nosematidae (Brooks, 1974).

Apparently species of Ataenis and Aphodius have not been examined for these parasites. A few investigators have reported nosematids from other Scarabaeidae. Steinhaus (1949) described Pleistophora (actually, Nosema) melolonthae

(Steinhaus) from the hemolymph of M. melolontha. Hurpin (1968) obtained the various parasitic stages of N. melolonthae from the fat body of M. melolontha. Nosematids have been found in a variety of coleopteran host tissues: Weiser (1961) found

Nosema curvidentis Weiser in the connective tissue of larvae and adults of Pitvokteines curvidens Germar (Scolytidae). Cali and Briggs (1967) described Nosema tracheophila Cali and

Briggs from the tracheal epithelium of Chrvsomela septempunctata (L.) (Chrysomelidae). Drea, et al. (1969) discovered a Nosema sp. in the gonadal tissues of alfalfa weevil, Hvpera postica (Gyllenhal) (Curculionidae). They

found that 90% of one laboratory colony of H. postica was

infected with this parasite and infected beetles exhibited

reduced egg and spermatozoa production. Weiser (1970) found

Nosema dendroctoni Weiser in the Malpighian tubules and muscle

of D. pseudotsuqae. Streett, et al. (1975) isolated a Nosema

sp. from the gut wall and lumen of Pissodes strobi (Peck)

(Curculionidae). They found the parasite in 3 0.4% of the

larvae and 68.1% of the adults examined. No external symptoms

of infection were apparent.

Coccidians (Protozoa: Sporozoa: Telosporea) are

intracellular parasites of both vertebrates and invertebrates

(Barnes, 1968). Nearly all of the coccidian parasites of

Coleoptera belong to the genus Adelina (Eucoccida: Adeleidae)

(Brooks, 1974). No coccidians have been reported from the

Aphodiinae,* however, Kawanishi, et al. (1974) reported that all of the A. spretulus examined for Bacillus spores also contained stages of an unidentified sporozoan in the fat body.

Weiser and Beard (1959) found Adelina sericesthis Beard in the fat body and subcuticular connective tissue of larvae of

Sericesthis pruinosa Dalman (Melolonthinae). They observed that larvae infected by this parasite appeared mottled brown, the gut being visible through the infected, translucent fat body. Hurpin (1968) isolated Adelina melolonthae Hurpin from the fat body of M. melolontha. 24

Gregarine Parasites

Gregarines (Protozoa: Sporozoa: Telosporea) are among

the most common parasites of invertebrates (Brooks 1974).

Kamm (1916) conducted one of the earliest and most

exhaustive studies on the gregarine parasites of insects. She

explained that the free-living adult stage, also called

trophozoite, sporont or sporadin stage, is most frequently

located in the mid-intestine of the host. It is sometimes

found in the pyloric caeca, and occasionally in the host body

cavity. Kamm observed that gregarines are generally host

specific. She reported Actinocephalus (Pvxinia) gregarines

(Eugregarinida: Actinocephalidae) from the intestines of

Aphodius prodromus Brahm, Aphodius nitidulus Fabricius, and

Rhizotroqus sp.. These parasites were also found in species

of three other genera of soil-inhabiting beetles: Carabus

(Carabidae), Staphvlinus (Staphylinidae), and Silpha (Silphidae).

Berberet and Helms (1969) reported two eugregarine parasites

from Phvllophaaa anxia (LeConte) (Melolonthinae): Greoarina

sp. (Actinocephalidae).

Nematode Parasites

Most of the nematode parasites of Scarabaeidae belong to

the genera Psammamermis (Mermithidae) , Bradvnema (Sphaerulari- dae), and Neoaplectana (Steinernematidae or Neoaplectanidae)

(Nickle 1974). Glaser and Fox (1930) and Glaser and Farrell

(1935) found the namatode Neoaplectana alaseri Steiner caused

40% mortality in larvae and pupae of P. Iaponica. Grubs 25

infected with this nematode were "flaccid and of an ocherous

brown tint". Nickle (1974) listed Neoaplectana hoptha Turco as

a parasite of P. iaponica, and N. melolontha Weiser, N.

georqica Kakulia and Veremchuk, Psammomermis korsakowi

Polozhentsev, and P. kulaaini Polozhentsev as parasites of

June beetles (Melolontha).

Insect Parasites

Several studies have been conducted on insect parasitism

of Scarabaeidae. Ritcher (1958) wrote that the insect parasites

attacking larvae of Melolonthinae, Rutelinae and Dynastinae

include many species of Tachinidae and Bombyliidae (Diptera),

and one species each of Pelecinidae and Ichneumonidae

(Hymenoptera). A few species of Pyrgotidae and Sarcophagidae

(Diptera) are known to parasitize adult scarabaeids. Tashiro, et al. (1969) reported parasitism of R. maialis larvae by

species of Dexilla, Microphthalma (Tachinidae), and Tiphia

(Tiphiidae).

Parasitic Fungi

Ritcher (1958) noted that the green muscardine disease,

Metarrhyzium anisopliae (Metchnikoff) Sorokin (Deuteromycetes or Hyphomycetes), is an important factor in natural mortality of white grubs in several continents, accounting for moderate to high mortality of Geotroous sp. (Melolonthinae), Cotinis texana Casey (Cetoninae), Euetheola ruqiceps (LeConte)

(Dynastinae), and Orvctes sp. (Dynastinae). Ritcher (1958) also wrote that Beauveria densa (Link) Picard (Deuteromycetes 26

or Hyphoraycetes) is parasitic on Melolontha larvae and that

species of Cordvceps (Ascomycetes or Pyrenomycetes) are a

sporadic parasite of scarabaeid larvae. Hurpin (1968) stated

that Beauveria tenella (Delacroix) Siemaszko was the most

frequently found pathogen in test populations of M. melolontha

at Versailles, France. Tashiro, et al. (1969) occasionally

found moribund R. maialis infected by M. anisopliae in field

plots.

A great deal of research has been devoted to the modes

of attack employed by the various parasitic fungi, as well as

to the environmental conditions under which parasitism is

most likely to occur. McLaughlin (1962) found that the white muscardine disease, Beauveria bassiana (Balsamo) Vuillemin, was most pathogenic to larvae, pupae, and adults of the boll weevil, Anthonomus qrandis Boheman (Curculionidae), under

conditions of high humidity. Schaerffenberg (1964) observed that M. anisopliae is thermophilic, growing most effectively at temperatures between 10 and 30 °C, whereas B. bassiana is not temperature dependent and can grow at temperatures ranging

from 0-40 °c. Schaerffenberg also observed that presence of lipid enhanced germination of both species. Walstad, et al.

(1970) found that, in general, both M. anisopliae and B. bassiana germinated and grew best under conditions of high humidity (92.5%) and high temperature (15 - 35 °C), on a non- sterile substrate. Hedlund and Pass (1968) observed that a test population of H. postica, infected with J3. bassiana, was 27

attacked by penetration of the sclerotized integument, resulting

in 100% mortality in 7-13 days. Gabriel (1968) determined that

B. bassiana, M. anisopliae, and Cordvceps militaris (L.) Link were all capable of producing chitinase and proteolytic enzymes enabling these fungi to penetrate the insect host's cuticle.

Insect Predators

According to Ritcher (1958) major insect predators of scarabaeids are certain larval Asilidae (Diptera), Tabanidae

(Diptera), Elateridae (Coleoptera), Histeridae (Coleoptera), and adult Carabidae (Coleoptera) and Formicidae (Hymenotptera).

Certain asilids have been observed to prey on Phvllophaaa and Anomala larvae; some tabanids are known to prey on

Phvllophaaa and Cvclocephala larvae. Carabids have been observed feeding on eggs and larvae of Rhizotroqus and Orvctes; various elaterids are known to feed on larvae of Orvctes,

Aphodius. and Melolontha. Certain histerids are reported predators of larvae of Aphodius, Teuchestes (Aphodiinae),

Oxythvres (Cetoninae), and Cetonia (Cetoninae); a number of formicids attack eggs and young larvae of scarabaeids.

Tashiro, et al. (1969) reported Harpalus pennsylvanicus Dejean

(Carabidae) as a predagor of R. maialis larvae, and Harpalus erraticus Say as a predator of R. maii1is eggs.

Vertebrate Predators

Ritcher (1958) wrote that to some extent, various reptiles and amphibians contribute to mortaility of scarabaeids by feeding on the eggs and larvae. He listed a number of birds, 28 including crows (Corvus), starlings (Sternus vulgaris L.), grackles (Quiscalus) song sparrows (Melospiza melodia

{Wilson)), meadowlarks (Sternella), woodpeckers (Piciformes:

Picidae), catbirds (Dumetella carolinensis L.), thrushes

Passerformes: Turdinae), robins (Erithacus rubecula L.), pheasants (Phasianus), gulls (Charadriiformes: Laridae), and owls (Strigiformes), that have been observed to feed on larvae of soil-inhabiting scarabaeids. Among the important mammalian predators Ritcher mentioned are moles (Insectivora: Talpidae), shrews (Insectivora: Soricidae), ground squirrels (Citellus), field mice (Rodentia: Cricetinae), raccoons (Procvon lotor

L.), and skunks (Carnivora: Mephitinae). Tashiro, et al.

(1969) reported starlings and skunks as the major predators of

R. maialis larvae in test plots of grub-infested turfgrass in

New York.

Phoretic Acari

Mites (Acarina) are sometimes found clinging to adult scarabaeids in the field. Costa (1969) explained that the simplest association between mite and host is pure phoresy, in which the mite is carried by its more mobile host to a specific habitat optimal for both host and mite. Usually the phoretic stage is only one stage in the life cycle of the mite. The type of phoretic stadium associated with the adult insect seems to be characteristic at the generic or even the family level. For example, Costa found only mature female mites of the genus Coleolaelaps on adult phytophagous scarabaeids; mature male and immature mites were found with the grub in its 29

subterranean cell, where they fed harmlessly upon the

exudates of the grub.

Costa (1969) observed two gasamine families of mesostig-

matic mites, the Dermanyssidae (17 species) and Pachylaela-

pidae (1 species), associated with phytophagous scarabs. He

reported five families of gasamine mites, the Eviphiidae (36

species, and (j> deutonymphs) , Macrochelidae (118 species,

deutonymphs), and Rhodacaridae (3 species), associated with

coprophagus scarabs. Evans and Hyatt (1963) reported 4

species of mites (Mesostigmata: Macrochelidae) associated

with coprid beetles (Coprinae). They collected Macrocheles

floridans Evans and Hyatt from Deltochilum orbiculare

Lansberge, Macrocheles scapularis Evans and Hyatt on Deltochilum

lobipes Bates, and Macrocheles hirsutissima (Berlese) on

Copris bituberculatus Cristofori and Jan and Dichotomius

carolinus (L.).

PHENOLOGY

Tashiro and Gambrell (1963) reported that observations of

the seasonal development of R. maialis over the years have

revealed consistent correlation between the occurrence of given

life stages and the flowering period of several common plants.

Three years of recorded correlations revealed that plant

development was accelerated or depressed in relation to temper­

ature in the same manner of R. maialis development. They

found these phenologica] observations helpful in predicting the

development of the various stages of the chafer, thereby eliminating the necessity of making soil surveys at frequent 30 and that by selective instead of random sampling he could at least double the number of egg pods found.

Salt and Hollick (1946) working with wireworms Agriotes sputator (L.) and A. obscurus (L.) and Burrage and Gyrisco

(1954) with R. ma1alis, observed changes in distribution with the age of the population. Egg clusters of these species were randomly distributed. Although the eggs and young larvae were clumped, the dispersion of the larvae tended to become random, or approached random, in later instars.

Finney (1941), working the 3 species of Agriotes wireworms, and Davis and Wadley (1949) with M. mexicanus egg pods, observed apparent changes in distribution resulting from changes in population density (i.e., the random distribution of low populations and the contagious distribution of higher ones). They explained that in a very sparse population of a typically clumped species, the chances of individuals occurring in any sampling unit is so low that their distribution is effectively random.

Southwood (1966) reported that true contagion in a population

may arise from patchiness of the habitat, including differential predation...or from the behavior of the themselves, or a combination of both. The behavior leading to aggregation in the absence of special attractive areas in the habitat, may be of two types; inter-individual attraction or the laying of eggs (or young) in groups... . III. GENERAL METHODS AND MATERIALS

The Cincinnati (CCC), Hyde Park (HPCC), Kenwood (KCC), and Losantiville (LCC) country clubs in Hamilton County, Ohio and the Terrace Park Country Club (TPCC) in Clermont County,

Ohio served as study sites for one or more aspects of the

3-year project. All 5 courses were within 13 km of each other and had histories of A. spretulus infestations on greens, tees and fairways. A turf laboratory was established at CCC to serve as the base of operations for the project from April,

1976 to November, 1978.

Parasitic and phoretic organisms removed from A. spretulus adults and larvae collected at HPCC, TPCC, and LCC during 1976-

1977 were identified. Observations on predators and phenology were made at all courses during 1977-78. Data on seasonal occurrence of adults in light traps, sticky traps, and soil samples were collected at TPCC, LCC, and KCC during 1977.

The golf course superintendent at each of these courses volunteered areas of two of his most heavily infested fairways for sampling and suggested secure areas for placement of adult- monitoring devices. Devices for monitoring adult activity were retained at KCC and TPCC in 1978. Sampling for life stages was restricted to TPCC. A study of overwintering

31 32 survivorship of beetles was conducted at TPCC, LCC, and KCC during 1977-78.

Soil samples were taken in overwintering sites at the wooded perimeters of the 3 courses (Appendix 1). Samples contained 1-4% organic material, 43-72% sand, 14-26% silt and

13-28% clay. The pH of the soil ranged from 5.8 to 7.3.

Bentgrass (Aqrostis species)and annual bluegrass (P. annua) were the primary grasses on fairways at these courses. Thatch accumulation on fairways was 0.6 - 1 cm at TPCC, 2-2.5 cm at

LCC, and 0.6-1.3 cm at KCC. Thatch accumulation on greens and tees was less than or equal to that on fairways at all 3 courses. Standard turf management practices, exclusive of insecticides (Appendix 2), were continued on the designated sampling areas at all 3 courses to maintain optimum turfgrass vigor. IV. INDIVIDUAL STUDIES

LIFE HISTORY

METHODS AND MATERIALS

A. spretulus eggs, larvae, pupae, and adults were observed systematically under field conditions. Measurements of 20 field- collected specimens of each life stage were obtained using a dissecting microscope fitted with an ocular micrometer.

Intestines of larvae and adults were dissected and examined for content.

Results

Dimensions of the life stages of A. spretulus, based on measurements of 20 specimens per life stage, are given in

Table 1. Eggs (Fig. 1) averaged 0.72 mm (0.67-0.76) long x

0.52 mm (0.42-0.55) wide. They were deposited in clusters of

11 or 12 within a cavity formed by the female in the lower 5-10 mm of thatch and upper 6 mm of soil among grass roots.

Occasionally eggs were found that were only 0.5-0.7 times as wide as typical field-collected eggs. When incubated at 2 3+

2 °C on moist filter paper the narrow eggs increased in diameter to the mean size in 48 hr*

First instar larvae (Fig. 2) averaged 2.44 mm (1.89-

3.78) long with head capsules 0.5 mm (0.46-0.55) wide. First

33 Table 1. Dimensions of A. soretulus life stages, based on measurements (in mm) of 20 field-collected specimens per life stage.

Length Width Head Capsule Width Life Stage Range Mean j S.D. Range Mean ± S.D. Range Mean ± S.D.

2gg 0.67-0.76 0.72+0.03 0.42-0.55 0.52+0.05 -

Instar 1 1.89-3.78 2.44±0.57 - - 0.46-0.55 0.5 + 0.03

Instar 2 3-78-6.09 4.75*0.68 - - 0.76-0.38 0.33±0.03

Instar 3 6.5-10.0 8.5 ±1.06 - - 1.22-1.34 1.3 +0.04

Pupa 4.2 -5-7 4.7 ±0.54 2.2 -2.6 2.5 ±0.14 -

Adult 4.0 -5.6 4.9 +0.41 1.8 -2.5 2.2 ±0.18 -

LO Fig. 1. Egg cluster of A. spretulus.

Fig. 2. First, second, and third instar A. spretulus. 36 instars were often found together, apparently in the vicinity of the original egg cluster.

Second instars (Fig. 2) averaged 4.75 mm (3.78-6.09) long with head capsules 0.83 mm (0.76-0.88) wide. They were found in the lower 5-10 mm of thatch and at the thatch-soil inter­ face and were not as aggregated as first instars.

Third instars (Fig. 2) averaged 8.5 mm (6.5-10.0) long with head capsules 1.3 mm (1.22-1.34) wide. Rasters were invested with teges of 40 to 45 irregularly placed, hamate setae

(Figs. 3 and 4). Mandibles had one long seta and one short seta dorsally (Fig. 5) and one seta ventrally. Third instars were most common at the thatch-soil interface. They were more dispersed than second instars.

Some larvae were found in thatch; others occupied cells formed 1 to 8 cm deep in the soil. Intestines of dissected specimens contained fine particles of roots and humus.

Pupae (Fig. 6) averaged 4.7 mm (4.2-5.7) long x 2.5 mm

(2.2-2.6) wide. They occupied cells excavated by the larval stage 1 to 8 cm deep in the soil.

Adults (Fig. 7) averaged 4.9 mm (4.0-5.6) long x 2.2 mm

(1.8-2.5) wide. Mature (fully sclerotized) adults were usually found in.the thatch. However, beetles were observed on turf- grass on warm, sunny, spring and fall afternoons, on summer evenings, and just prior to and following summer rainfall.

Teneral (red-reddish brown) adults were observed in pupation cells, tunneling in soil, and at the thatch-soil interface.

Intestines of dissected specimens contained particles of humus. Fig. 3. Rastral pattern of mature A. spretulus larva.

0.5 mm Fig. 4. Scanning electron micrograph of hamate setae on raster of mature A. spretulus larva. 38

Pig. 5 • Dorsal aspect of left larval mandible of A. spretulus. showing long seta (a) and short seta Tb). Fig. 6. Female (a) and male (t>) A. spretulus pupae Arrow indicates aedeagal protuberance of male.

Fig. 7. A. spretulus adult. 40

At least 100 observations of ovaries in dissected females

revealed that individual ovarioles, of which there were 12/

female, contained 3-4 eggs in different stages of development.

At least 50 females that were captured and dissected in

late May and June, 1978, exhibited evidence of having already oviposited. In each case proximal ends of ovarioles were con­ stricted and vacant where mature eggs were located prior to oviposition. Near-proximal and median regions of ovarioles contained ova in moderate and early stages of development, respectively. One female captured in June, 1977 contained one mature egg at the proximal end of one ovariole, whereas the other ovarioles contained much smaller ova at their proximal ends. Apparently this female had deposited only 11 of its 12 mature eggs. The number of times that a female oviposits could not be determined.

DISCUSSION

A comparison of my findings with those of other researchers concerning the seasonal history of the life stages of A. spretulus and certain other turfgrass scarabaeids may serve to aid in the correct identification of A. spretulus in the future.

The occasional discovery of thin eggs capable of enlarging under moist conditions suggests they had been ovi­ posited shortly before sampling and had not yet absorbed much water from the soil. These observations support those of

Ludwig (1932) who wrote that eggs of P. iaponica and other 41

scarabaeids must absorb water prior to and during embryonic

development to survive.

A. spretulus eggs can be distinguished from those of P.

japonica and R. maialis on the basis of size. Newly deposited

P. japonica eggs measure 1.5 x 1-1.5 mm (Fleming, 1972), making

them at least 2x larger than A. spretulus eggs. Newly

deposited R. maialis eggs measure 0.73 x 0.49 mm (Tashiro, et al.

1969). These dimensions are similar to those of A. spretulus eggs that are 48+ hr old. A. spretulus eggs are deposited in

clusters in the thatch and upper 6 mm, whereas R. maialis eggs are found singly and deeper in the soil. Eggs of Aphodius qranarius (L.), an occasional pest of golf course turfgrasses

(Jerath & Ritcher, 1959; Niemczyk, 19 78; Niemczyk & Power, 19 78) have not been described.

Weaver and Hacker (1978) reported that the average length of a third instar A. spretulus is 0.25 in. (6.4 mm). The smallest third instar I measured was 6.5 mm long. Hoffmann

(19 35) and I agree on the average length of the third instar larva, 8.5 mm. However, his observation of 0.75 mm as the average width of the head capsule is less than my observation of 1.22 mm as the minimum width. The head capsule widths of

Jerath's (1960) 3 unknown Ataenius spp. larvae ranged from 1.19-

1.35 mm. Based on these measurements, any one of these 3 larvae could have been A. spretulus.

The larvae of A. spretulus, P. japonica, and R. maialis are similar only when third instars of the former are compared to first instars of the latter two species. Upon close inspection, P. iaponica and R. maialis larvae are much more 42

setose and possess more complex rastral patterns (Fleming,

1972; Tashiro, 1969) than those of A. spretulus. The distinc­

tion between larvae is not as apparent between A. spretulus and

A. oranarius. Equivalent instars are similar in size. However, the raster of A. oranarius possesses a palidium, or V-shaped arrangement of spines (Jerath, 1960, Niemczyk, 1978), while that of A. spretulus does not.

Hoffman (19 35) and I found the average length of pupae to be 4.7 mm. His reported average pupal width, 2.1 mm, was less than mine, 2.5 mm. Weaver and Hacker (1978) stated that pupal measurements were about the same as those of adults. However,

I found adults to be 0.1-0.2 mm longer and narrower than the pupae from which they eclosed.

A. spretulus pupae can be sexed by the method of Tashiro, et al. (1969) for R. maialis and Fleming (1972) for P. japonica.

Males possess a protuberance on the 10th abdominal sternum, whereas females do not.

All investigators reported similar length and width ranges for A. spretulus adults. A. spretulus adults are easily dis­ tinguished from those of most other scarabaeid species infesting turfgrass, except A. qranarius. However, A. oranarius adults possess transverse carinae on the meso- and metatibiae

(Woodruff, 1973) ; whereas A. spretulus adults do not.

My observations of adults in flight indicated that A. spretulus is a weak flyer, traveling only short distances at estimated speeds of 1-3 m/sec at altitudes of 0.3-4.6 m in still air. Flying beetles settled quickly with each wind gust. 43

Fleming (1972) reported that most flights by P. japonica cover short distances, but that beetles are capable of traveling

2 to 15 mi (3.2-24 km) per year, averaging 5 mi (8 km).

Fleming observed little flying on cool windy days.

Tashiro, et al. (1969) observed that R. maialis may attain altitudes of 60 to 80 ft (18.3-24.4 m) and distances up to 725 yd (663.4 m) in sustained flight. Maximum annual dispersal was estimated to be 4 to 5 mi (6.4-8 km).

The oviposition behavior of A. spretulus is conspicuously different from that of P. japonica and R. maialis. An insemin­ ated A. spretulus female burrows 1 cm into the soil and deposits all the mature eggs she contains in one cluster. Although no field-collected female oviposited more than one cluster of 12 eggs in the laboratory, individual females seem capable of ovipositing more than one egg cluster apiece.

According to Fleming (1972) an impregnated P. japonica female burrows 5-10 cm into the soil, deposits 3 or more eggs, one to a cavity, and emerges to feed and copulate again. A female may enter the soil 16 or more times and deposit up to

133 eggs (typically, 40-60) during her life.

Tashiro, et al. (1969) reported that during its lifetime an inseminated R. maialis female oviposits 2-46 eggs (av. 22) , singly, 5-10 cm deep in the soil.

SAMPLING FOR LIFE STAGES

METHODS AND MATERIALS

Sampling sites (Fig. 8) were marked out in unrestricted random sample design. Each site was 30.5 x 18.3 m and contained

15 equal-sized square plots 6.1 m on a side. The corners of Green or Tee End of Fairway-

■18.3 m 1 2 3

4 5 6

7 8 9 e vr\ Rough Rough o

10 11 12

13 i: 14 15 a i 1i b

Fig. 8. Typical 558.15 m sampling site, sub-^ divided into plots (a) and quadrats (b), in which unrestricted random sampling for life stages of A. spretulus was conducted. 45

each plot were marked with aluminum-capped wooden stakes driven

flush with the ground. Turf around stakes were periodically

sprayed with Paraquat ® to prevent overgrowth. Each plot was

further subdivided into quadrats 3.05 m on a side and identi­

fied by plot number (1, 2, ...,15) and position letters (T =

tee end, G = green end, R = right side, L = left side).

Unrestricted random sample placements were generated by drawing marked plastic chips from a jar. Each of 60 chips was painted with a different plot number and quadrat designation.

Chips were thoroughly mixed in the jar before each drawing.

Drawn chips were returned to the jar before subsequent drawings.

Samples were taken with a 10.8 cm i.d. golf-hole borer to a depth of 7-8 cm. Each sample was examined in the field for

10-15 min.; numbers of all A. spretulus life stages were 2 2 recorded and coverted to No./ft (per 929 cm ) by multiplying

No./sample x 10.15. Examined turf samples were replaced and watered to facilitate regrowth. Watered samples usually filled in with new turf within 2-3 weeks. Previously sampled turf plugs were subsequently avoided.

Life stage sampling was conducted on TPCC fairways 14 and

17, LCC fairways 11 and 16, and KCC fairways 4 and 6 in 1977.

Each location was sampled twice per week, with 10 samples per site on each occasion.

Sampling was intensified at and restricted to two sites on

TPCC fairways 14 and 17 in 1978. Plots were sampled over a two 46

day period each week, taking 20 samples per site. On May 20,

1978 the TPCC staff inadvertently treated the sampling site on

fairway 17 with insecticide, rendering it unsuitable for

further sampling. A second sampling site was established at

the end of fairway 14, opposite the original sampling site on that fairway. The original sampling site was then designated

14A and the new area 14B.

All adults and pupae obtained were sexed and the resulting data tested for significant deviation from a 1:1 sex ratio using the normal approximation to a binomial distribution.

Some adult females obtained each month were examined to indicate the percentage inseminated.

RESULTS

Data obtained from life stage sampling at TPCC, LCC and

KCC during 1977 are given in Appendices 3, 4, and 5, respectively.

Sampling data gathered from TPCC during 19 78 are listed in

Appendix 6. Dated, major events in the development of A. spretulus, as observed in sampling, are summarized in Table 2.

The period required for A. spretulus to pass through one generation was 65^ 5 days at soil temperatures of 25 - 6 °C

(range = 13-38 ° c ) , based on first generation periods estimated from life stage sampling data (Figs. 9 and 10). First genera­ tion periods were determined as the number of days between appearances of first and second generation eggs in samples.

A suspicious increase in the duration of the second genera­ tion, coupled with depressed population (age class) peak 47

Table 2. Summary of A. spretulus life stage sampling record at TPC( life Stage fivent 1977 TPCC do. per No. per tlo. per No. per Date Sample 929 cm2 Date Sample 929 cm2 Adults Initially found...... 9/19^ - 3 9/15®^ - 2 Maximum no. foundt In one sample...... 7/8 13 (*132) 7/15 19 (*193) As mean of 20 samples.•• 7/8 - 37 7/15 - 27 No. found on last sampling date* Maximum... 10/18 3 (^ 30) 10/20 2 (* 20) Mean..... iO/I8 - 6 10/20 - 7 Teneral adults Initially Pound...... 6/20 - - 6/30 --

*05" 5/3 6 5/2 15 Maximum no. foundt In one sample...... 8/11 2* (*299) 5/9 39 (*395) As maan of 20 samples.•• 5/10 - 31 5/9 - 39 9/7 -- 8/29 -- Larvae 5/10 - - 5/16 - - and 5/20 - - 5/23 - - Prepupae Third instars initially found...... 5/27 -- 5/31 -- Prepupae Initially found...... 6/7 ■ - - 6/8 - - Maximum no. of larvae and prepupae found! 6/10 39 (*396) 6/16 62 (*629) As mean of 20 samples... 6/10 127 6/16 • 125 No. of larvae and prepupae found on last sampling datei Maximum...... 10/18 5 <* 51) 10/20 3 (* 30) Mean...... 10/18 - 19 10/20 - 9

Pupae Initially found...... 6/1 it _ (* 13) 6/21 _ (* 6) Maximum no, foundt In one sample...... 6/21 16 (*162) 7/5 29 (*299) As mean of 20 samples.•. 6/29 - 98 7/7 - 35 No. found on last sampling datei Maximum... 10/18 1 (* 10) 10/20 3 (* 30) Mean..... 10/18 9 10/20 5

g / First sampling date at given golf course. 47

g-pretulus life stage sampling record at TPCC, LCC, and KCC during 1977 - 1978.

1977 1970 TPCC LdS "" KCC TPCC too. per too. per too. per No. per No. per No. per No. per No. per Date Sample 929 cm2 Date Sample 929 cm2 Date Sample 929 cm2 Date Sample 929 cm2 Jj/H jS/ - 3 4/15*^ - 2 4/18 - 0.5 4/5-68'' - 0.5 7/8 13 (*132) 7/15 19 (*193) 8/30 9 <* 91) 7/12-13 14 (*142) 20 samples. e 7/8 - 37 7/15 - 27 9/6 - 21 7/12-13 - 31 et Naxlmum. • 10/18 3 30) 10/20 2 (* 2 0) 10/18 2 (* 20) 10/25 3 (* 30) Mean.... 10/18 - 6 10/20 - 7 10/18 - 4 10/25 - 5 6 /2 0 - - 6/30 - - 6/27 -- 7/5 --

5/3 _ 6 5/2 - 15 5/9 _ 5 5/24 _ 37 pie...... 8/1 1 24 (-244) 5/9 34 (*345) 8/15 20 (*203) 7 /2 6 24 (*244) ' 20 samples. • 5/18 - 31 5/9 - 34 7/19.8/18 - 18 8/ 9-10 - 40 ••4e«e*4*4*o « 9/7 -- 8/29 - - 8/30 -- 9/14 -- MtMttiMU • 5/10 - - 5/16 -- 6 /2 -- 5/31-6/1 -- It i i • t m u1 » fr 5 /2 0 -- 5/23 -- 5/31 -- 6/ 7 -8 -- Mtdtiitttt • 5/27 -- 5/31 - - 5/31 - - 6/ 7-8 - - 8/7 . - - 6 /8 - - 6/9 -- 6/ 21-22 -- tupae foundt 6 /1 0 39 (*396) 6 /1 6 62 (*629) 6/13 40 (*4o6) 8/9-10 35 (*355) V 20 samples. 6 /1 0 - 127 6/16 - 125 8 /1 2 - 99 6/ 21-22 - 124 md on last 10/18 5 (* 51) 10/20 3 (* 30) 10/18 3 (* 30) 10/25 1 (* 10 ) 10/18 - 14 10/20 - 4 10/18 - 3 10/25 - 1

6 /1 (* 13) 6/21 _ (* 6) 6 /2 0 _ (* 2) 6/28-29 - (« 64) iple...... 6/2 1 16 (* 162) 7/5 24 (*244) 7/12 20 (*203) 7/6 20 (*203) ' 20 samples. • 6/24 - 48 7/7 - 35 8/25 - 24 7/5-6 - 50

:et Maximum. e 10/18 1 (* 10) 10/20 3 (* 30) 10/18 3 (* 30) 10/25 1 (* 10) Mean.... • 10/18 9 10/20 5 10/18 5 10/25 1

sn golf course. 48 densities at TPCC and LCC (Figs. 9 and 10) were greatly due to a long period of oviposition by first generation females during

July through September. This relationship of increasing ovi­ position periods to successive generations of multivoltine populations was described by Clark, et al. (1967). The second generation was not used to estimate generation period for this reason.

An apparent increase in peak density of the second genera­ tion over that of the first was observed at KCC. This was probably due to oviposition by extant first generation females in the sparsely-populated sampling sites during mid-summer.

The sex ratios of 754 adults and 698 pupae obtained from

TPCC (Appendices 7 and 10), 469 adults and 558 pupae from LCC

(Appendices 8 and 11), and 530 adults and 507 pupae from KCC

(Appendices 9 and 12) during 1977 generally were not signifi­ cantly different (P > .05) from 1:1. The same was true of the sex ratios of 1080 adults and 634 pupae collected at TPCC during

1978 (Appendices 13 and 14). However females outnumbered males in 67% of the monthly sex ratios prepared from data collected both years.

The inseminated fraction on a monthly basis, of 248 females collected at TPCC, LCC, and KCC during 1977 was 100% in April and May, 77 - 25% in June, 43 - 39% in July, 82 - 11% in August, and 76 - 3% in September. The inseminated portion on a per- month basis, of 276 females examined from TPCC in 1978 was 100% in April, May and June, 5 2% in July, and 76 i 3% in August,

September and October. CM No./ 929 cm Instar 1 30 Instar Instar Instar Instar 302 Instar ua 30 Pupae n Adults 30 Adults i. * esnl curneadaudne fA srtls ie tgs in stages life spretulus A. of 1977-1978. abundance during and TPCC at sites occurrence sampling Seasonal 9* Pig. 3 Uo Uo 20 Uo 50 1,0 80 50 50

P , A . U . U . U , E . OCT . SEP , AUG . JUL . JUN MAY . , APR aD6^SEP_rOCT

CM No./ 929 cm Instar Instar Instar 2 30 2 Instar ntr 130 Instar ua 30 Pupae dls 30 Adults i. 0 Saoa ocrec n bnac o . peuu ie tgs in stages life spretulus A. of abundance and occurrence Seasonal 10. Fig. sampling sites at LCC and KCC during 1977. during KCC and LCC at sites sampling 3 P , A t JUN MAY t , APR PT 1VIAY APFTT U SP OCT SEP AUG

O Ui 51

DISCUSSION

The sampling method used was selective to the extent that sampling sites were established in locations with a history of infestation. However, samples were taken from each site (Fig. 8) in unrestricted random fashion. The selective location of the sampling sites assured a large population density from which to draw samples. The random sampling program served to disclose the type of distribution in the chosen site.

The golf-hole borer produced sample units of sufficient 2 size (91.6 cm ) to provide a reasonable balance between the proportion of the population being sampled and the time necessary for examining the sample. However, the nature of the sample unit did not lend itself to both expedient in-field processing and accuracy in counting the life stages therein.

This fact is evident in the discrepancy between numbers of eggs, first instars and second instars found in samples during May and

June (Appendices 3-6) and numbers of third instars and pupae found in the same areas during June and July. Figs. 9 and 10 also show that the peak number of eggs found in samples is substantially less than the peak number of larvae. However, it is possible that numbers of larvae surpassed numbers of eggs in samples if an equilibrium existed between the number of eggs being deposited and the number hatching in the sampling sites.

The evidence for two generations per year was strong at the

3 courses sampled. Increased numbers of adults, including tenerals, collected in life stage samples in late June and early July, 1977 and 1978 indicated the completion of the first 52 generation (Appendices 3-6; Table 2? Figs. 9 and 10). An abrupt increase in the number of eggs found in samples around mid-July, which followed a gradual decline, during June and early July, signaled the beginning of a second generation

(Appendices 3-6; Figs. 9 and 10). Similar evidence for two generations per year was reported by Niemczyk and Dunbar

(1976) in Ohio and Connecticut and by Weaver and Hacker (1978) in West Virginia.

In 1977 the peak, mean numbers of first and second gener- 2 ation third instars in TPCC samples were 87 and 48 per 929 cm , respectively (Figure 9). In 1978, corresponding first and 2 second generation peaks were 112 and 26 per 929 cm , respectively.

Comparison between first generations from 1977 to 1978 suggests there was an increase in population size from one year to the next. However, comparison between second generations suggest there was a decrease in population density in a year's time. the same trends were reflected in numbers of all life stages found in TPCC samples over the two-season span (Figure 9).

If peak numbers of first and second generation third instars are averaged for each year, the average for 1977, 2 (87 + 48)/2 = 67.5 per 929 cm , is very close to the average for 2 1978, (112+26/2 = 69 per 929 cm . This suggests that A. spretulus population at TPCC, as monitored from sampling sites, increased very little, if at all, from 1977 to 1978.

An extended second oviposition period (see Clark, et al.,

1967) was greatly responsible for the apparent decrease in peak infestation density of the second generation in TPCC 53

sampling sites both years (Figure 9) and in LCC sites in 1977

(Figure 10). However, this trend was not observed in KCC

sampling sites during 1977. Instead, numbers of third instars

and other life stages of the second generation exceded those

of the first (Figure 10) .

Apart from the amplitude-reducing effect of lengthened

oviposition period on plotted data, the numerically-reduced

second generation infestations in sampling sites at TPCC and

LCC could have resulted from one or more density-dependent

factors. Diminished food in sampling sites that were densely-

infested by the first generation may have initiated dispersal of first generation adults. Perhaps predators reduced numbers of first generation adults in sampling sites, resulting in a smaller egg cohort in the second generation. Finally, parasites may have increased during the first generation and then dampened the second generation.

One density-independent factor that may have reduced the second generation in TPCC and LCC sampling sites was the effect of insecticide on the surrounding population. All greens, tees, and fairways, except sampling sites, were treated with insecticides 2-3 times per year at both golf courses. Such treatments would not necessarily affect dispersal from sampling sites but could have reduced numbers of incoming adults from adjacent turfgrass.

The large second generation infestation in KCC sampling sites probably resulted from oviposition by extant first generation females and migrant females from adjacent turfgrass. 54 individuals. Possibly, most of the females found during

August through October consisted of a few inseminated first generation individuals and second generation females that copulated prior to seeking overwintering sites. However, many of the females collected in July were probably newly- eclosed and had not yet contacted males. It was impossible to determine the period required for sexes to locate each other in the field following eclosion. No mating was observed in samples or elsewhere. In contrast, adults of P. japonica have frequently been observed in. copula. Males approach virgin females as they emerge from the soil ca. 9 days after eclosing (Fleming, 1972).

GEOGRAPHICAL SIZE OF POPULATION

METHODS AND MATERIALS

For statistical analysis, observations from sampling sites were organized by age class (egg, instar 1, instar 2, instar 3, prepupa, pupa, adult) and by number of samples con­ taining a given number of individuals of a given age class

(Xq samples contained 0 adults, x^ samples contained 1 adult, x- samples contained 2 adults, ..., x samples contained y ^ y adults).

Contingency tables were used to determine whether sampling sites at a given course contained distinct A. spretulus populations or members of one population. In these tests, the greater the probability (P) value, the more liekly observations from compared sampling sites belonged to the same population.

A 1% level of significance (P > .01) was used due to the small 55 sample size, as dictated by the time-cost factor of the project.

RESULTS

The first contingency table comparison was made between infestations in TPCC 14 and 17, using 1977 data (Table 3).

Significant similarities were evident between these infesta­ tions in 5 out of 7 age classes: eggs, first instars, prepupae, pupae, and adults.

A second comparison was conducted to validate the re­ establishment of the second sampling site at TPCC from fairway 17, which had been inadvertently treated with insecti­ cide, to fairway 14, on the end opposite from sampling site

14 (i.e. changing 14 to 14A and 17 to 14B) in May, 1978. The validating comparison between 17 and 14B could not be made using 1978 data because 17 had to be abandoned before enough observations could be recorded to conduct the test. Therefore, a comparison was made between 1977 data from 17 and 1978 data from 14B (Table 4). Similarities were significant in 4 out of

7 age classes first instars, prepupae, pupae, and adults. The similarities were sufficient to support substitution of 14B for 17 in 1978.

A third comparison was made between 1977 data from 14 and

1978 data from 14A (Table 5) to test for infestation similarities in the same sampling site from year to year. Similarities were significant in 4 out of 7 age classes: eggs, and instars 1,

2, and 3. 56

Table 3. X, comparison of A. snretulus infestations in TPCC sampling sites 14 and 17 by age class, using 1977 data.

Age Class X 2 D.P. P > .01

Eggs 3.86 4 .425 *

Instar 1 5.48 3 .140 *

Instar 2 22.36 7 .002 n. s.

Instar 3 38.94 13 .000 n.s. * O 5.28 • 00 Prepupae 6 Vn

Pupae 10.14 8 • 255 *

Adults 11.74 7 .109 * 57

Table 4. X comparison of A. soretulus infestations in TPCC sampling sites 17 (1977 data) and l4B (1978 data) by age class.

Age Class X2 D.F. P > .01

Eggs 11.46 3 .009 n. s. Instar 1 7.01 3 .072 * Instar 2 22.81 7 .002 n. s. Instar 3 48.19 13 .000 n.s. CO H Prepupae 7.49 5 • * Pupae 6.98 7 .431 * Adults 15.13 7 .034 *

Table 5- 7? comnarison of A. snretulus infestations in TPCC sampling sites 14 (1977 data) and 14A (1978 data) by age class.

Age Class X2 D.P. P > .01

Eggs 2.82 3 .420 * Instar 1 2.90 3 .407 # Instar 2 15-40 7 .031 * Instar 3 12.61 10 .246 * 0 0 Prepupae 18.72 6 • n.s. Pupae 23.11 7 .002 n.s. Adults 34.03 6 .000 n.s. 58

A fourth comparison was made between observations from

14A and 14B (Table 6) to test for infestation similarities in

the same fairway. Similarities in this test were significant

in all age classes.

The final infestation comparison was made among pooled

1977 observations from TPCC, LCC, and KCC (Table 7) to test

whether the infestations at the 3 courses belonged to the same

population or were distinct populations. Similarities were

significant in only 2 out of 7 age classes: eggs and pupae.

DISCUSSION

Within-course comparisons of observations from sampling

sites at TPCC (Tables 3-6) suggest that infestations on the

same fairway and on separate fairways at the same course were

not significantly different (P >.01) and, therefore, probably belonged to the same population.

A between-course comparison of sampling observations from

TPCC, LCC, and KCC (Table 7) indicated that infestations at the sites on the 3 courses were distinct statistically, though not necessarily different biological populations.

DISPERSION PATTERN OF POPULATION

METHODS AND MATERIALS

Two models were used to determine the theoretical distribution which best described the A. spretulus dispersion pattern in fairways at the 3 courses. Observations were tested for random distribution by the Poisson model and for clustered (contagious) distribution by the negative binomial model. The Chi-square test was used to compare the observed 59

ft Table 6. % comparison of A. snretulus infestations in TPCC sampling sites 14a and 14b by age class, using 1978 data.

Age Class 7? D.F. P > .01

Eggs 1.86 3 .602 * Instar 1 8.0 5 3 .045 * Instar 2 15-41 6 .017 * Instar 3 14.91 10 • 135 * Prepupae 9.66 5 .085 * Pupae 15.14 7 .034 * Adults 4.75 6 • 576 *

ft Table 7. X comparison of A. snretulus infestations in sampling sites at TPCC, LCC, and KCC by age class, using 1977 data pooled by course.

Age Class X2 D.F. P > .01

Eggs 25.29 12 .014 * Instar 1 48.73 10 .000 n.s. Instar 2 47.35 20 .000 n.s. Instar 3 83.44 30 .000 n.s. Prepupae 45.45 14 .000 n.s. Pupae 34.24 18 .012 * Adults 55-79 14 .000 n.s. 60 frequency distributions of numbers of each age class/sample with expected frequencies for the Poisson and negative binomial distributions.

The formula (Southwood, 1978) used for calculating probabilities for the Poisson model was -x -X £- P (x) = e x i where x = the number of individuals of a given age class/sample

X = the sample mean, and e = the base of natural (Napierian) logarithms 2.72)..., and x! may be obtained from tables of 2 factorials. The associated Chi-square (x ) had two fewer degrees of freedom than the number of comparisons that were made between observed and expected frequencies. Frequencies with small expectations (< 5) were pooled.

The formula (Southwood, 1978) used for calculating the probabilities for the negative binomial model was

I-1 (x + TO... f x ) X ( k \ k P(x) = xi r (1c) + k / \ X + k / , where x, x, and x! are as explained earlier, (P) = log gamma functions obtainable through tables, and k = the dispersion parameter, with iterative formula:

N loge (l A-) - 2 , where N = the total number of samples, logg = Napierian logarithms, and Ax = the sum of all frequencies of sampling units containing 2 more than x individuals. The x associated with the negative binomial model had 3 fewer degrees of freedom than the number 61

of comparisons that were made between expected and actual

frequencies.

Ten samples/site were taken twice per week during 1977.

When data were fit to the two distributions, the negative 3 binomial described the dispersion pattern better. The

method of Rojas (1964) was used to suggest the desired number

of samples/site/occasion to be taken during 1978, to maintain

a low standard error. The recommended number of samples ranged

from 263 - 4480/site/occasion (for a standard error of 10%),

depending on the age class to be sampled. The recommended

number of samples for eggs and young larvae was about 6x/that

for third instars through adults, to achieve the same precision

(ca. 90%) in sampling.

A suggested sampling program based on the method of Rojas

(1964) was not practical because of the tremendous time-

cost factor involved and resulting irreparable damage to the

fairways and the impact on the beetle population. Instead, the number of samples was doubled in 19 78 at TPCC and sampling

at LCC and KCC was discontinued. Therefore, the 20 samples/ site/wk program was adopted at the expense of precision,

resulting in a standard error >_ 36%.

RESULTS

Observations on A. spretulus infestations in TPCC 14 and

17 made during 1977 were fit independently, by age class and fairway, to the Poisson and negative binomial models to test for population distribution differences between sampling sites at the same course. 62

TPCC 14 and 17 observations, fit to the Poisson model,

exhibited no significant similarities between observed and

expected frequencies in any age class. However, these same

data, when fit to the negative binomial model, exhibited

significant similarities in 6 out of 7 age classes: first

instars through adults (Tables 8 and 9) .

Pooled 1977 observations from TPCC 14 and 17, when fit

to the Poisson and negative binomial models, gave the same

results (Table 10) as those obtained when observations from

TPCC 14 and 17 were fit to the models individually.

Pooled 1977 observations from LCC 11 and 16 and from KCC

4 and 6, when fit to the Poisson model, exhibited no similarity between observed and expected frequencies in any age class.

However, observations from both locations, when fit to the negative binomial model, exhibited significant similarities in

5 out of 7 age classes (Tables 11 and 12).

DISCUSSION

Data presented in tables 16-20 indicate the distribution of A. spretulus in fairways at the 3 courses sampled is better 2 described by the negative binomial model for contagion (S

2 - 2 — (s >x, 2) the Poisson model for randomness (s = x) .

However, the consistent fractional values, 0.0150 - 0.4 395, of the dispersion parameter, k, of the negative binomial fit of all populations and age classes sampled, suggest that the dispersion pattern may be described most accurately by a 2 _ logarithmic distribution (S > x, 1> k > 0). Table 8. Fit of A. snretulus population from TPCC 14 to Poisson and negative "binomial distributions by age class, using 1977 data. Poisson Negative Binomial Age Class X2 D.F. P > .01 X2 D.F. P>,.01

Eggs 338.1 2 .000 n.s. 25*66 2 .000 n.s. Instar 1 96.4 1 .000 n.s. 3*98 1 .046 • Instar 2 264.3 2 .000 n.s. 1.6 4 .809 * Instar 3 809.9 4 .000 n.s. 15*9 10 .103 * Prepupae 210.7 2 .000 n.s. 3*22 4 .522 * Pupae 276.5 2 .000 n.s. 5.27 6 .510 * Adults 90.2 2 .000 n.s. 3-31 4 .507 *

Table 9» Fit of A. spretulus population from TPCC 17 to Poisson and negative binomial distributions by age class, using 1977 data. Poisson Negative Binomial Age Class X2 D.F. P>.01 X2 D.F. P>.01

Eggs 376.7 2 .000 n.s. 23.4 3 .000 n.s. Instar 1 367*2 2 .000 n.s. 3-35 2 .187 # Instar 2 491.8 3 .000 n.s. 9.7 6 .138 * Instar 3 1892.5 6 .000 n.s. 20.1 11 .044 * OO O Os Prepupae 158.9 2 .000 n.s. 1.6 4 • * Pupae 200.6 2 .000 n.s. 6.3 5 .278 * Adults 241.7 3 .000 n.s. 5.3 5 .380 * 64

Table 10. Fit of A. snretulus population at TPCC (pooled 1977 data from sampling sites 14 and 17) to Poisson and negative binomial distributions by age class. Poisson Negative Binomial Age Class X2 D.F. P >.01 X2 D.F. P>.01

Eggs 933. **6 3 .000 n. s. 162.68 6 .000 n.s. Instar 1 149.86 1 .000 n.s. 4.52 3 .211 * Instar 2 848.52 3 .000 n.s. 9.82 8 .278 * Instar 3 2997-81 7 .000 n.s. 29.17 15 .015 * Prepupae 260.19 2 .000 n.s. 3-53 6 .740 * Pupae 576.00 3 .000 n. s. 11.97 7 .102 * Adults 458.77 3 .000 n.s. 10.26 5 .068 *

Table 11. Fit of A. suretulu s population at LCC (pooled 1977 data from sampling sites 11 and 16 ) to Poisson and negative binomial distributions by age class. Poisson Negative Binomial Age Class X2 D.F. P >.01 X2 D.F. P > .01

Eggs 1072.81 3 .000 n.s. 83.35 6 .000 n.s. Instar 1 473-22 2 .000 n.s. 23-60 5 .000 n.s. Instar 2 881.76 3 .000 n.s. 21.67 9 .010 * Instar 3 2814.39 6 .000 n.s. 21.29 15 .128 * Prepupae 298.86 2 .000 n.s. 8.70 5 .122 * Pupae 536.71 3 .000 n.s. 5-98 7 .542 * Adults 275-58 2 .000 n.s. 4.21 5 .520 *

Table 12. Fit of A. snretulus population at KCC (pooled 1977 data from sampling sites 4 and 6) to Poisson and negative binomial distributions by age class. Poisson Negative Binomial Age Class X2 D.F. P > .01 X2 D.F. P > .01

Eggs 227.05 1 .000 n.s. 57.44 4 .000 n.s. Instar 1 57.81 1 .000 n.s. 0.23 1 .632 * Instar 2 351.59 2 .000 n.s. 3.84 5 • 573 * Instar 3 1982.19 5 .000 n.s. 41.02 13 .000 n.s. Prepupae 111.15 1 .000 n.s. 3.50 3 .321 *

O * Pupae .000 • 0 377.52 3 n.s. 12.60 6 In Adults 329.81 2 .000 n.s. 8.38 4 .079 * 65

It is unclear whether the dispersion pattern of A.

spretulus in fairways was dictated by effect of patchiness of

the habitat on oviposition habits of females (see Landin 1961,

1968 concerning Aphodius, and Handford, 1955, concerning M.

mexicanus egg pods), the laying of eggs in clusters (see

Burrage & Gyrisco 1954 concerning R. maialis. Guppy & Harcourt

1970 concerning Phyllophaaa. and Salt & Hollick 1946 concerning

Acriotes), conspecific attraction (see Fleming 1972 concerning

P. iaponica, Vite & Pitman 1968, Wood, et al. 1970 concerning

Ips and Dendroctonus), or a combination of these factors

(Southwood 1966) .

The nature of A. spretulus infestations on golf courses

poses many sampling and management difficulties. A sequential

sampling program for this insect based on the negative binomial

distribution cannot easily be prepared due to the inherent

small k value and high probability of incorrectly assessing

population density (Type I and II errors).

A trial sequential sampling program for A. spretulus was

constructed according to procedures described by Ives and

Warren (1965) and Southwood (1978): 2 Hq : _< 50 3rd instars/929 cm sample 2 H^: >_ 100 3rd instars/929 cm sample

k (3rd instars) = 0.32

Unacceptably large areas of indecision between plotted accept­

ance and rejection lines resulted when values of 0.05 and 0.10 66 were used as probabilities of incorrect assessment. Reason­

able areas of indecision could be obtained only when error

probabilities of 0.48 or 0.49 were used:

d Q = 69.33n + 25.13

d1 = 69.33n - 25.13, where a = 3 = 0.48

d Q = 69.33n + 12.56

= 69.33n - 12.56, where (X = 3 = 0.49

However, a sequential sampling program based on error probabil­

ities of 48-49% would be of questionable value. Ives and

Warren (1965) were able to establish a sequential sampling

program for Phvllophaqa larvae; however, they worked with a

k value of 1.5936, as opposed to the 0.0150 - 0.4395 k range

associated with A. spretulus infestations.

FLIGHT AND INSEMINATION STUDIES

METHODS AND MATERIALS

Rectangular screen sticky traps and 8-vaned hardware

cloth sticky traps were evaluated for capturing A. spretulus

adults in 1977 and 1978 respectively. Rectangular traps were

based on the "sticky screen" traps of Prescott and Newton (1963)

and consisted of prefabricated aluminum window screens supported by steel rods driven into the ground (Fig. 11). Traps were

located approximately 1.2 m above ground, had a screen mesh 2 2 size of 2.2 mm , a mean capture area of 0.24 m , and were

coated with Tack Trap ®. A second coat was applied to traps in

July to maintain adhesiveness t w o rectangular traps per course, 67

Fig. 11. Rectangular, screen sticky trap used to monitor seasonal flight activity of A. spretulus in 1977.

Fig. 12. Eight-vaned sticky trap used to monitor seasonal flight activity of A. spretulus in 1978- 68 were located and monitored at TPCC, LCC, and KCC from April 12 through October 31, 1977.

Traps were redesigned for 1978 to study vertical distri­ bution of flying adults and to increase capture/unit time.

The eight-vaned traps used in 1978 were based on the baffle design of Coon and Rinicks (1962) and the multi-level design of Taft and Jernigan (1964). They consisted of squares of hardware cloth bolted at right angles to one another, in two groups of 4, on wooden posts driven into the ground. Vanes were arranged at right angles to permit capture of flying adults approaching from any direction. The groups of vanes were set at approximately 0.91 and 1.82 m above ground (Fig. 12).

Vanes were grouped at two heights to test for possible seasonal differences in beetle flight density at different altitudes. 2 2 Each trap had a capture area of 0.74 m , a mesh size of 40 mm , and was coated with Mapco Stikum Second and third coats were applied to traps in June and August to maintain adhesiveness. The adhesive surface of the hardward cloth was considered less likely to deflect air movement and wind-borne adults than the adhesive-coated screens used in 1977. Four

8-vaned traps at TPCC and two at KCC were monitored daily from

March 13 through October 31, 1978.

Sticky traps were positioned 30-300 m apart in wooded rough and near wooded golf course perimeters. Traps were adjacent to life stage sampling sites and suspected overwintering sites. Captured adults were counted and removed daily to define onset and duration of immigration from overwintering 69

sites to fairways in spring and onset and duration of emigra­ tion from fairways to overwintering sites in the fall. Sticky

traps were examined in the A.M. or early P.M. to avoid

combining data from two days. Each month, from May through

October, 1977, an average of 36 females, selected at random

from traps at all 3 courses, were examined for insemination by checking spermathecae for sperm.

Light traps were positioned at the borders of infested fairways to provide a daily record of beetle flight and activity throughout the season. Live adult females were dissected to determine reproductive status at biweekly intervals.

Two types of suction light traps were used, author-made traps in 1977 and commercially-made traps in 1977 and 1978.

Both trap types utilized unfiltered U.V. lamps and vacuum fans, and were fitted with 10 mm mesh excluder screens to keep out large insects. Author-made traps (Fig. 13) were discontinued during 1978 because they lacked the functional convenience and operational dependability of the commercially made Will-o'-the- (r ) Wisp Buglite Fish Feeder w traps, fitted with funnel nets and collection jars (Fig. 14) .

One author-made trap at TPCC, and one commerically-made trap per course at LCC and KCC were monitored from 28 March through 31 October 1977. Two commercially-made traps at TPCC and one at KCC were monitored from March 14 through 31 October

1978. All traps were mounted 2-3 m above ground and operated continuously. Captured insects were removed and counted during the A.M. or early P.M., daily. Excluder screens, fan 70

Fig. 13. Author-made suction black light trap used in 1977 to monitor daily and seasonal A. spretulus flight activity.

Fig. 1*K Modified Will-o'-the-Wisp Buglite Fish Feeder® suction black light trap used in 1977 and 1978 to monitor daily and seasonal A. spretulus flight activity. 71 parts, and funnel screens were cleaned of insect parts several

times both years.

The normal approximation of a binomial distribution was used to determine if the proportion of inseminated females varied between months during 1977 and 1978.

RESULTS

Appendices 15-18 list all sticky trap capture data from

TPCC, LCC and KCC during 1977. The largest captures of beetles

(1-18/929 cm of trap surface area/day) on sticky traps during

1977 occurred in April and July through October (Fig. 15). 2 Adult captures were rare (0-0.6/929 cm /day) during May and

June.

All of the females examined in May, 67% in June, 69% in

July, 96% in August, 91% in September, and 94% in October, were inseminated.

Appendices 19-21 list all 1978 sticky trap capture data from TPCC and KCC. 2 The largest captures of adults (1-16/929 cm /day) at TPCC and KCC during 1978 occurred in April, May, and August through 2 October (Fig. 16). Only a few adults (0 - 0.6/929 cm /day) were captured in June and July.

Significantly more (P> .05) beetles were found on the lower vanes (0.91 m) than on upper vanes (1.83 m) during

April, May, and October (Table 13). Fig. 15. Seasonal occurrence and abundance of A. suretulus adults on rectangular screen sticky traps (averaged per day) at TPCC, IjCC, and KCC during 1977.

13.17 9.33 6.511 5.90

111 J i l l . llll. JiL III.lln. in, ji ill 1 ill In II APR MAY TT.KIN JUL AUG SEP “ OCTI

Fig. 16. Seasonal occurrence and abundance of A. spretulus adults on R-vanod sticky traps (averaged per day) at TPCC and KCC during 1978. Table 13. Captures of A. spretulus on 8-vaned sticky traps at two heights in 1978. Significance between captures at different levels and relationship to average monthly temperature are given.

(6 ft) level Number 0.91 m (3 ft) level 1.83 m lil Capture Ratio Average Month Examined Between Levels Air No./level Percent of total No./level Percent of total (P >.05) Temperature

12.6 APR 2273 1221* 53.85 101*9 1*6.15 .0002 * 16.7 MAY 530 288 5^.3^ 21*2 1*5.66 .01*56 * 22.8 JUN 13 10 76.92 3 23.08 .0521* n.s. 23.9 JUL 1*5 19 U 2.22 26 57-78 .2981* n.s. 23.2 AUG 363 175 1*8.21 188 51.79 .1*966 n.s. 22.0 SEP 297 15^ 51.85 11*3 1*8.15 .5222 n.s. 11.8 OCT 572 320 55.91* 252 1*1* T 06 .001*6 *

Total **093 2190 53.51 1903 1*6.1*9

Appendices 22-25 list light trap data from TPCC, LCC and

KCC during 1977. The largest captures of beetles (80-837)/trap/ day) in light traps during 1977 occurred in April and from

July through October (Fig. 17). Few adults (0-38/trap/day) were captured in March, May, and June.

The inseminated fraction of 288 females examined from light traps during 1977 was significantly greater (P> .05) than the virgin fraction for every month except July (Table 14).

Appendices 26-28 list all light trap data from TPCC and

KCC during 1978. The largest captures of adults (40-390/ trap/day) during 1978 occurred from May through October

(Fig. 18). Few adults (0-19/trap/day) were captured during

March and April.

The inseminated fraction of 273 females collected in light traps during 1978 was significantly greater (P> .05) than the virgin fraction in every month except July (Table 15).

DISCUSSION

Results of the sticky trap and light trap studies in 19 77 and 1978 (Figs. 15 and 16) indicate that greatest flight activity over the rough near sampling sites and at the golf course perimeters occurred in Spring, late Summer, and Fall. Since traps were located in surroundings like those reported by

Hoffmann (1935) and Weaver and Hacker (1978) as harboring over­ wintering adults, these early and late season captures undoubt­ edly consisted of adults migrating to and from overwintering sites. Capture height data from 8-vaned traps (Table 13) further revealed that adults generally flew lower to the ground

(<1.83 m) near traps at these times than in early and mid-summer. No./Trap 100 Uo 20 60 70H 80 90 h 30-I 50-| H Fig. 17. ..Seasonal occurrence and abundance of A. spretulus adults in author-inade in Will-o*- and adults spretulus A. of abundance and 17. occurrence ..SeasonalFig. leWs® uto lc lgt rp (vrgdprdy a PC I3, n C drn 1977- IX3C, during TPCC, at KCC day) and per (averaged traps light black suction tlie-Wisp® uto bak ih tas aeae per- 1978. (averaged at. traps during day) KCC light and TPCC black suction i. 8 Saoa ocrec adaudne fA sntls dls nW11-o'-the-Wisp® Wi1 in adults sinetnlus A. of abundance and occurrence Seasonal 18.Fig. Record No A APR MAR iuL 109 LiL T MAY + ± JUN l L jj . I JU1, AUG Jg'l SEP 155

OCT

76

Table 14. Percentage of inseminated A. snretulus females captured in black light traps at TPCC, LCC, and KCC during 1977*

Deviation from No. 29 Inseminated Month 1:1 Ratio of Examined Inseminated to Percent No Percent Yes Virgin 92 (P<.05)

MAY 32 31 96.88 1 3.12 .0000 *

JUN 60 54 90.00 6 10.00 .0000 *

JUL 75 35 46.67 40 53.33 .5620 n . s .

AUG 52 50 96.15 2 3-85 .0000 *

SEP 40 38 95.00 2 5.00 .0000 *

OCT __2 100.00 _0 0 .0026 *

Total 263 217 81.94 51 18.06

Table 15• Percentage of inseminated A. soretulus females captured in black light traps tat TPCC and KCC during 1978.

Deviation from No. 22 Inseminated Month 1:1 Ratio of Examined Inseminated to Percent No Percent Yes Virgin 22 (P<.0 5 )

MAY 56 56 100.00 0 0 .0000 *

JUN 44 44 100.00 0 0 .0000 ♦

JUL 54 30 55-56 24 44.44 .4122 n.s.

AUG 82 75 91.46 7 8 .54 .0000 * * SEP _J7 96 .49 ii, 2%. .0000 Total 273 237 86.81 36 13.19 77

Table 16. Effect of site, month, and sex on survivorship of overwintering A. spretulus adults near Cincinnati, Ohio.

Dependent Variable! Ratio Sum of Mean Sourpg Squares Sapors F Value P > T R-Souare C,V. Model 3? 5-4596 0.1^76 32.94 0.0001 0.9870 36.2127 Error 16 0.071? 0.001*5 Std. Dev. Rat jo Mear. Corrected Total 53 5.5313 0.066Q 0.1645 Spupcg D.F. Tvue 1 SS F Value F > F Site 2 0.0594 6.63 0.0080 * Month 6 4.e550 135.48 0.0001 * Sex 1 0.0102 2.26 0.1503 n.s. « Site/Month 16 0.3795 5-29 0.0009 Site/Sex 2 0.0176 1.95 0.1702 n.s. Month/Sex e 0.1377 3.84 0.0105 *

Table 17. Effect of site and month on survivorship of overwintering A. spretulus adults near Cincinnati, Ohio.

Dependent Variable! Ratio Sum of Mean Spyrpg D.F. Squares Souare F Value P> F R-Souare C.V. Model 26 5.2938 0.2036 23.16 0.0001 0.9571 50.7329 Error 27 0.2374 0.0088 Std. Dgv. Ratio Mean Corrected Total 53 5.5312 0.0936 0.1648 Source fiJL. Tvoe 1 SS F Value P > F Site 2 0.0594 3.36 0.0491 n.s. Month 6 4.6550 69.03 0.0001 * Site/Month 16 0.3795 2.7C 0.0111 n.s. 78

This behavior suggests either inability to achieve higher

altitude or closeness of point of debarkation to destination

(overwintering site to oviposition site; emergence site to overwintering site).

The few adults captured in light and sticky traps in

June represent the last adults to leave overwintering sites and those seeking second or third oviposition sites at other locations. Although no field-collected females deposited more than one cluster of 12 eggs in capativity, evidence from dissections indicated that females, after ovipositing once, relocated by flight to oviposit again.

July trap captures corresponded to the completion of the first generation as described by Weaver and Hacker (1978) for bivoltine populations of A. spretulus in West Virginia.

First generation adults may have been captured while seeking mates or suitable oviposition sites.

Early spring and fall captures in light traps were usually not proportionately as large.as those in sticky traps due to low nocturnal temperatures at these times. Light trap captures during June and July were proportionately larger than sticky trap captures. The attractive nature of the U.V. lamps versus the nonattractive nature of the sticky traps probably was responsible for this difference.

Two suction black light traps were built in 1977; there was insufficient time to build more. Testing revealed a faulty electrical system in one trap which proved irreparable. When the immediate need for 3 traps became apparent, only two commercially-made traps were available for use. Therefore, 79 one of my traps had to be used in addition to the two modified

Will-o'-the-Wispl^ devices in 1977. device was obtained for use in 1978. In general, the commer­ cial devices were simpler and more dependable to operate than my own.

Light traps and sticky traps were both important adult monitoring systems, although the latter were useful in more ways. Light traps were more effective qualitatively, as activity indicators due to their attractiveness, although useful only on warm evenings. Sticky traps were more useful in detect­ ing diurnal activity on warm or sunlit days with cool evenings.

Strategically placed sticky traps also were reliable indicators of migratory periods in the insect's seasonal history. Both systems were equally useful as sources of data on sex ratio and percentage of females inseminated.

OVERWINTERING STUDIES

METHODS AND MATERIALS

In March and April, 1977, over 100 6-7 cm-deep samples were taken with a golf-hole borer at suspected A. spretulus overwintering sites at TPCC, LCC, and KCC and examined for presence of beetles. No specific data were recorded; however, results of this general survey led to formulation of a sampling program to characterize overwintering sites in terms of adult survivorship at these golf courses. 2 Five samples, 929 cm x 5 cm deep, were taken monthly from suspected overwintering sites at TPCC, LCC, and KCC from

September, 1977 through May, 1978. Samples consisting of 80 soil and duff were taken from the edge of wooded areas (Figs.

19 and 20) in the rough and perimeters of golf courses.

Samples were taken in the same 4 x 3 m area at each course to minimize differences between samples.

Samples were placed individually in plastic bags and transferred indoors. Frozen samples were allowed to thaw before examination. The soil was carefully broken apart and hand-sifted into a 19 1. container half filled with water.

All floating organic debris was skimmed from the water with a fine screen scoop. Drained flotation material was processed in Berlese-Tullgren funnels to extract adults. Numbers of living and dead adults and living inseminated females obtained from Berlese samples and from residual debris on funnel screens were recorded.

Efficiency of the flotation method was defined by pro­ cessing 10 samples and then inspecting the sediment. Overall,

735 beetles were floated from 10 samples and the sediment contained 8 beetles. Therefore, the flotation method was estimated to be [735/(735 + 8)] x 100 = 98.92% efficient.

RESULTS

The first series of samples taken during the general survey of overwintering sites indicated increasing population density from the fairway to the edge of adjacent wooded areas and decreasing density from the tree line into the wooded areas.

Few beetles were found in turf samples taken in the rough where no trees were found. Beetle numbers increased markedly in 81

Pig. 19. Typical overwintering site of A. snretulus adults at perimeter of golf course.

Fig. 20. A. snretulus adult overwintering site in rough between fairways at a golf course. 82 samples taken within 1 to 2 m of the edge of wooded areas.

The greatest density of overwintering beetles was found in samples taken within a 2 to 3 m wide swath at the tree line.

Samples taken farther than 2 to 3 m into a wooded area con­ tained fewer beetles.

The tree line of wooded areas surrounding golf courses was often characterized by the presence of tall fescue grass

(Festuca arundinacea Schreber), herbaceous or woody ground cover such as Lamiurn amplexicaule L. and Toxicodendron radicans

(L.) Kuntze, border weeds such as Impatiens species and

Polvmnia canadensis (L.), and duff accumulations. Beetles were most abundant on the soil beneath duff and in the top

1 to 5 cm of soil. Immature stages were not found at these sites. 2 2 Maximum beetle density observed was 264/ft (or 929 cm ) at KCC in October, 197 7. Approximately 6 5% of all overwintering beetles found were female and 89.7% of those examined were inseminated.

Beetle survivorship declined somewhat from September through March at all 3 courses. Low survivorship ratios in

April and May reflected the absence of vigorous adults which may have already migrated back to golf course turf. If the

March survivorship rate is taken as representative of cumulative winter survival, then from 90 to 96% of the 1977 populations at the 3 courses survived to 19 78. 83

Population density data were tabulated by site, month, sex, and mean survivorship (Appendix 30). Influence of the first 3 variables and their interactions on beetle survivor­ ship were tested with a 3-way analysis of variance. The results (Table 16) indicated that the effect of sex on sur­ vivorship was not significant (P< .05), suggesting that males and females overwintered with approximately equal success.

A 2-way ANOVA was performed using the same data with the sex and sex interaction effects deleted from the model. The results (Table 17) indicated that the site and site/month interaction effects were not significant (P< .01) to the survivorship of overwintering adults under the conditions described. The month effect was the greatest of the 3 factors with an F value of 69.03. The site and site/month effects had

F values of 3.38 and 2.70, respectively. The inclusiveness of the 2-way model was reflected in an R-square value of

0.96. However, the importance of the month effect was revealed in an R-square value of 0.88 when tested in a one-way ANOVA.

DISCUSSION

The rate of winter survivorship, 90-96%, of A. spretulus was high compared to rates of certain other Coleoptera that overwinter near the soil surface, such as 85% survival of

Dendroctonus ponderosae Hopkins (Watson 1971) and 72% survival of Dendroctonus rufipennis (Kirby) (Frye et al. 1974). However, the overwintering behavior and survivorship of A. spretulus was very similar to that of Coleomeqilla maculata (DeGeer) as reported by Benton and Crump (1979). 84

Male and female A. spretulus overwintered with equal

success. Apparently this is not the case with all Coleoptera.

Watson (1971) and Safranyik (1976) observed that female D. ponderosae survived cold and winter better than males.

The site effect reflected the suitability of specific overwintering sites, including nature of soil and type and quantity of cover vegetation or duff at the 3 courses. The cumulative survival at the 3 sites did not vary significantly

(P> .05), 95.5+0.23%, from September, 1977 through March, 1978.

No consistent effect on survivorship could be attributed to a specific soil composition of pH at the sampling sites.

Apparently a pH range of 5.8-7.3 and soil composition ranges of

1-4% organic matter, 43-72% sand, 14-26% silt, and 13-28% clay constituted suitable overwintering conditions for A. spretulus.

CORRELATION OF PLANT PHENOLOGY. TEMPERATURE. AND HUMIDITY WITH

SEASONAL LIFE HISTORY OF A. spretulus

METHODS AND MATERIALS

Daily records of phenological events were made from March 24 to November 1, 1977 and 1978 at various locations in Hamilton and

Clermont counties. Key points in the annual cycle of A. spretulus were correlated with conspicuous, concurrent botanical events, such as onset of blossoming, seedhead appearance, and seed release of common woody ornamentals, weeds, and wildflowers.

Daily air temperature and precipitation data were obtained from the National Oceanic and Atmospheric Administration 85

recording station at the Abbe Observatory in Cincinnati, Ohio,

approximately 13 km from the farthest sampling station. Soil (r) temperature at 5 cm was monitored with a Weiss^ soil probe

thermometer during 19 77 and part of 1978 at TPCC, CCC, and KCC.

In 1978, soil temperature data were obtained with a Honeywell®

circular chart, 7-day recorder buried in a protective encase­

ment at the edge of KCC fairway 4 (Pig. 21). The recorder

probe was positioned 1.8 m horizontally into the fairway, 5 cm

deep.

Correlations of daily soil and air temperatures and

precipitation with daily light and sticky trap captures were

made in an effort to establish the threshold flight temperature

for A. spretulus and to elucidate effect of humidity on

flight activity.

RESULTS

Inability to obtain eggs from laboratory-reared A.

spretulus made it impossible to generate a threshold temper­

ature base for development. Therefore, a flight threshold was

substituted for the usual developmental threshold for using

day degree accumulations as a means of predicting seasonal

activity and development of A. spretulus in the field.

Validity of the flight threshold was based on the premise that propagation of a species cannot occur unless the minimum temperature necessary for the physical activity of every life

stage is reached. Flight is an important aspect of A. spretulus biology because it is the most effective means of moving between summer and winter habitats, bringing the sexes together, and locating suitable oviposition sites. Fig. 21. Protective encasement with circular chart, 7-day recorder used to record soil temperature 5 cm deep in a sampling site at KCC during 1978. The minimum temperature at which adults were captured

on sticky traps was between 13 and 15 °c. Captures occurred

on traps at TPCC on October 8 and 17 and at TPCC and KCC on

October 24, 1978 (Appendices 19 and 20) when daily maximum air

and soil temperatures were in this range. The developmental

threshold of A. spretulus may also lie in the 13-15 °c range

since Ludwig (1928) observed no embryonic development in eggs

of P. iaponica below 13 °c and no hatching below 15 °c.

Therefore, I selected 13 °C, the lower limit, as the flight

threshold to avoid the possibility of overestimating the

temperature at which adults of A. spretulus become physically

active.

Day degree accumulations (DDA) were calculated from the

first day the mean air temperature exceeded the flight

threshold to the last day life stages were sampled in fairways

(23 February - 31 August, 1977 ; 31 March -3 September, 1978).

Accumulations were calculated for each major point in the biology of A. spretulus in the field. The formula (adapted

from Arnold 1960) used for calculating day degree accumulations was :

DDA = I(daily mean air temperature - 13).

The DDA information relating to A. spretulus biology to temperature during 1977 and 1978 was combined with the phenological information (Table 18) to demonstrate the

synchronous relationship between temperature and plant and

insect development from year to year. The result was a method Table 18. Day degree basis of A. spretulus biology and associated plant phenology.

A. spretulus Phenology Day Degree Approximate Biological Plant Species Eroit Accumulations Time of Event Year

Onset of migration of Crocus 10 Early to late overwintered adults (Crocus veraus L.) March to fairways Appearance of first / Vanhoutte spirea YYSr^ 100 - 150 Early to mid- generation eggs " (Spiraea vanhouttei (Briot)Zabel) May Spreading cotoneaster FFB (Cotoneaster divaricata Rehder,Wilson) __ Horse chestnut FFB (Aesculus hippocastanum L.) Black cherry FFB (Prunus serotina Ehrhart) Black locust OB (Robinia pseudoacacia L.) Columbine OB (Acuilegia x hvbrida) Annual fleabane OB (l£i£S£OIi annuus (L.)Person) , Cottonwood 0SR-' (Populus deltoides Marsh)

Appearance of first Regal privet . OB 170 - 190 Mid- to late generation larvae (Lieustrum obtusifolium Siebert.Zuccarini) May Multiflora rose OB (Rosa multiflora Thunberg) Chinese peony OB (Paeonia lactiflora x hvbrida)

Predominance of Milkweed FFB 360 - 1*60 Mid- to late mature first (Asclepias svriaca L.) June generation larvae Adam's needle FFB and appearance of (Yucca filamentosa L.) first generation v/ Black-eyed Susan FFB pupae (Rudbeckia hirta L.) American basswood (Linden) FFB (£Uia americana L.) Queen Anne's lace (wild carrot) OB (Daucus carota L.) Staghom sumac OB (Rhus tvphina L.)

Appearance of first Plantain lily OB 470 - 560 Late June to generation adults (Hosta plantaginea x hvbrida) early July Summer phlox OB (Phlox paniculata L.) Black cherry FFB (Prunus serotina Ehrhart) Black locust OB (Robinia pseudoacacia L.) Columbine OB (Aouilegia x ljyb£ida) Annual fleabane OB (Eriaeron annuus (L.)Person) Cottonwood OSR^/ (Ponulus deltoides Marsh)

Appearance of first Regal privet OB 170 - 190 Mid- to late generati on larvae (Ligustrum obtusifolium Siebert.Zuccarini) May Multiflora rose OB (Rosa multiflora Thunberg) Chinese peony OB (Paeonia lactiflora x hvbrida)

Predominance of Milkweed FFB 360 - 460 Mid- to late mature first (Asclenias svriaca L.) June generation larvae Adam's needle FFB and appearance of (Yucca filamentosa L.) first generation b / Black-eyed Susan FFB pupae (Rudbeckia hirta L.) American basswood (Linden) FFB (£Uia americana L.) Queen Anne's lace (wild carrot) OB (Daucus carota L.) Staghorn sumac OB (Rhus tvnhina L.)

Appearance of first Plantain lily OB 470 - 560 Late June to generation adults (Hosta plantaginea x hvbrida) early July Summer phlox OB (Phlox paniculata L.) Appearance of second.. / Rose of Sharon OB 650 - 710 Early to late generation eggs (Hibiscus svriacus L.) July Woodland sunflower OB (Kelianthus strumosus L.)

Appearance of second Field thistle OB 740 - 800 Mid- to late generation larvae (Cirsium discolor (Muhlenberg)Sprengel) July

Predominance of mature second generation Tall ironweed OB 1000 - 1140 Early to mid- larvae and appearance (Vemonia gjgantea (Walter)Britton) August of second generation^ pupaen n n a o 3/

Appearance of second Canada goldenrod FFB 1100 - 1160 Late August to generation adults and (Solidago canadensis L.) early September onset of migration to overwintering sites a. Appropriate point in A* spretulus biology for use of adult control program (Niemczyk 1978). b. Appropriate point in A. spretulus biology for use of larval control program (Niemczyk 1978). c. Onset of blossom. d. First full bloom. e. Onset of seed release. 89 for predicting A. spretulus activity and development by moni­

toring temperature and/or concurrent botanical events.

The effect of temperature on development, as observed in

successive years, is demonstrated in Fig. 9. Average monthly

temperatures in the Cincinnati area were 1-6 °C warmer in

1977 than in 1978, during February, March, April, and May.

This difference between years in warming trend was reflected

in a 10-15 day lag in development of the TPCC population in

1978 behind that of 1977. This was especially evident in

first generations.

Precipitation appeared to have a flight-suppressing effect on A. spretulus adults when daytime temperatures were < 18 +

1 °C. However, there was a pronounced increase in afternoon and evening flight activity just prior to and during light precipitation at air temperatures > . 1 8 + 1 °c.

DISCUSSION

The plants most useful in predicting onset of spring oviposition by A. spretulus, namely Spiraea vanhouttei (Briot)

Zabel, Robinia pseudoacacia L., and Aesculus hippocastanum L., were among the important phenological indicators used by

Tashiro, et al. (1969) to predict seasonal development of R. maialis. These common, easily recognized plants may be of value in predicting seasonal life history events of other economically important insects,* thereby serving as indicators for timing preventive control strategies in situations where no visible damage to the hostplant can be tolerated. The timing of insecticide applications for turfgrass pests is especially 90

important in light of thatch penetration difficulties and

short residual times demonstrated for many organophosphates

(Niemczyk, et al. 1977; Sears & Chapman 1979).

The idea of calculating day degrees using a threshold

temperature based on an event other than development is not new. Apple (1952) based a day degree accumulation program for

following seasonal development of the European corn borer,

Ostrinia nubilalis (Hubner) on the planting threshold for

Zea mays L. rather than on the developmental threshold of the

insect.

An increase in flight activity before and during light precipitation on warm evenings may be typical of other aphodiine scarabs. Woodruff (1973) collected several adults of

Pleurophorus longulus Cartwright just before a light rain. He noted that beetles continued to fly in the rain. The same is not true of all scarabaeids. Fleming (1972) reported that P.

iaponica flight is retarded at relative humidity > 60% and ceases on rainy days.

ASSOCIATED ORGANISMS

METHODS AND MATERIALS

A record was kept of all predators, parasites, and phoretics thought to be associated with A. spretulus. Whenever possible, organisms found on and in adults and immatures were preserved for identification. 91

RESULTS

Five percent or less of larvae and pupae from life stage samples and those reared in the laboratory showed symptoms of disease. Body contents of the affected immatures consisted entirely of a light brown or black liquid. Ten discolored larvae and pupae were examined for presence of parasites. A few larvae contained abundant spores of a nosematid micro- sporidian. One larva contained immature stages of a neo- aplectanid nematode. No other parasites were detected in discolored insects.

Larvae at all 3 courses exhibited symptoms typical of infection by a milky spore bacterium (Fig. 22) in June, 1977.

Incidence of milky disease (Table 19) at all study sites increased from <1% in May and June to 25-29% in October.

Maximum levels of infection coincided with natural decline in larval density as adults were produced at the close of the second generation (Appendices 3-5). Milky disease incidence in larvae at TPCC in 1978 increased erratically from a minimum of 0% in May to a maximum of 7% in September. During both years, approximately 95% of the infected larvae were third in­ stars; the remaining 5% were second instars and prepupae.

One third instar larva collected at TPCC in August, 1977 contained 4 cephaline eugregarine sporonts (Fig. 23). The sporonts were aligned side by side, subdermally, in the hemocoel and were visible through the transparent cuticle and dermis layers in the anterior third of the larva, along the 92

Table 19. Incidence of milky spore-diseased A. spretulus at TPCC, LCC, and KCC during 1977 and 1978.

Percentage of Larvae and Prepupae Infected Month TPCC LCC KCC

1977 1978 1977 1977

MAY 0 0 0 0

JUN 0.47 0.16 0.55 0.15

JUL 6.67 5-34 9.39 2.00

AUG 7.39 3.71 2.88 0.20

SEP 18.31 6.79 10.56 1.74

OCT 27.27 4.76 28.57 25.00 22. Milky spore diseased larva of A. spretulus ompared to healthy larva (h).

1 mm

Fig. 2 3. Cephaline eugregarine sporont found in third instar larva of A. spretulus shape of living sporont (a) and photo­ micrograph of preserved specimen (b). 94 •

dorsal line. The sporonts measured approximately 1 mm long

by 0.2 mm wide. No further investigation was made of the host

larva for parasites.

Most moribund larvae, prepupae, pupae, and adults found

intact in life stage samples and rearing studies were covered with mycelium and spores of a Hyphomycetes fungus.

Three species of carabid beetles were found in life stage sampling sites at the 3 courses: Amara lecontei (Csiki),

Amphasia interstitialis (Say), and Harpalus fulqens Csiki.

A. lecontei was the most commonly encountered of these.

Carabids were not observed feeding on A. spretulus immatures but were suspected of having done so, based on reports by

Balduf (1935) and Ritcher (1958).

The most likely vertebrate predator of A. spretulus larvae was the starling, S. vulgaris. Small flocks were frequently observed pecking at infested fairways at all 3 courses. I suspect barn swallows (Hirundo rustica L.) may have preyed upon

A. spretulus adults in flight at KCC, based on frequent sight­ ings of these birds feeding just above fairways.

A. spretulus adults captured in flight or collected from turf in the spring and fall often had mites on them. Deutonymphs of the mesostigmatic family Uropodidae were often found, attached singly or in clusters by anal pedicels to the elytra and sterna of beetles collected at TPCC and HPCC in April and May,

1976. One deutonymph of Parasitus sp. (Mesostigmata:

Parasitidae) was found clinging to the head and pronotum of a beetle at HPCC in May, 19 76. Three deutonymphs of Dendrolaelaps 95

sp. (Mesostigmata: Digammasellidae) were found beneath the

elytra of all beetles captured in flight at LCC in October,

1976. An adult female of Macrocheles sp. (Mesostigmata:

Macrochelidae) was found clinging to the head and thorax of a

beetle collected at HPCC in May, 1977. Female Macrocheles

insignitus (Berlese) were found on thoracic sterna of beetles

collected in flight at LCC in May and October of 1976. Female

Scutacarus sp. (Prostigmata: Scutacaridae) were found on

thoracic sterna of beetles captured in flight at HPCC in May

and at LCC in October, 1976. Deutonymphs of Histiostoma sp.

(Astigmata: Anoetidae) were found on the thoracic and abdominal

sterna of beetles collected at LCC in August, 1976.

DISCUSSION

Natural enemies of A. spretulus were too few in number to consistently maintain population density at a non-damaging

level at TPCC, LCC, and KCC from 1976 through 1978. Conse­ quently, natural enemies alone cannot be considered adequate

in a management strategy for this insect on golf courses.

Microsporidian, gregarine and nematode parasites (and possible fungal parasites) of A. spretulus were found in only

a small fraction of the insects examined.

Incidence of milKy spore bacilli in fairways apparently was not high enough to provide acceptable control of A.

spretulus on many golf courses in the Cincinnati area. The effect on the more destructive first generation was apparently negligible. More second generation larvae were affected, but not enough to dampen the population satisfactorily during

1977-1978. 96

Birds, such as the starling and barn swallow, were

observed on all courses, but their consumption of A. spretulus

adults and larvae did not seem to cause a noticeable decline in

the number of life stages found during sampling, except

possibly at KCC. Regular, thorough insecticide treatment

programs were not practiced at KCC during 1977-1978, and birds

seemed most numerous there.

LABORATORY OVIPOSITION STUDIES

METHODS AND MATERIALS

Female beetles were collected from overwintering sites

and placed in 250 ml paper cups containing 8 different sub­

strate combinations (Table 20). Five replicates of 3-5

females/cup were used to test each substrate for its suit­

ability for oviposition. Replicates of 7 substrate types

were held from 19 to 30 days at 23+2 °c, 68+12% R.H., and 14 h

photoperiod. An illumination intensity of 885 + 334 lux was (K) supplied by Westinghouse Agro-Lite ^ fluorescent lamps. Cups were examined weekly for eggs. Replicates of the Pa-Th-Sl-Sd-

Gr substrate were held under similar conditions of humidity

and temperature but were subjected to natural daylight and were undisturbed for 2 7-32 days.

RESULTS

The only substrate in which females oviposited was the

Poa annua-thatch-soil-sand-gravel combination (Table 20). One

female deposited 12 eggs in one replicate and two females Table 20. Suitability of various substrates on egg maturation in and oviposition by inseminated, overwintered females of A. spretulus at 23±2 °C.

Replicates Substrate./ No. Test 1 2 3 ^ 5 Mixture * Per and Ratio Replicate (Days? E D ^ E C ^ S ED EO S ED EO S ED EO S ED EO 3E IE IE s 2E Sl-Sd-D M 2M 2M 2M 1M 84-15-1 3 30 3 I 0 3 L 0 3 L 0 3 1L 0 3 L 0

IE 2E 3E IE E C -Sl-Sd-D 2M 1M M 1M 2M 3O-54-15-I 3 30 3 L 0 3 L 0 3 L 0 3 1L 0 2 L 0

E E E E E M -Sl-Sd-D M M M M M 5O-34-15-I 3 19 0 L 0 0 L 0 0 L 0 0 L 0 0 L 0

E E E E E C -M -Sl-Sd M M M M M 10-50-30-10 3 19 0 L 0 0 L 0 0 L 0 0 L 0 0 L 0

E IE IE E IE C -3d-D 2M 1M 2M M 2M 60-39-1 3 19 3 1L 0 2 L 0 3 L 0 3 3L 0 3 L 0

E E E E & M -Sd-D M M M M M 60-39-1 3 19 0 L 0 0 L 0 0 L 0 0 L 0 0 L 0

E E E E & C -M -Sd-D M M M M M 24-60-15-1 3 19 0 L 0 0 L 0 0 L 0 0 L 0 0 L 0

E E E E £ M M M M M PS-Th-Sl-Sd-Gr^ 5 32 0 L 0 0 L 0 1 1L 12 2 2L 23 0 L 0

a. Substrate mixture legend; C = cow manure; D = diet supplement mixture of powdered milk, soy flour, and brewers yeast; M * Milorganite; Sd = sand; SI = soil. b. Female survivorship. c. Stage of egg development (i.e., S = early, M = moderate, L = late) per female. d. Number of eggs oviposited. e. Poa Annua (Pa) growing in thatch (Th) on consecutive layers of soil, sand, and gravel (Gr). 98 deposited single clusters of 11 and 12 eggs, respectively, in another. Larval hatch and survivorship was 16.7% in one replicate and 87.5% in the other at termination of the study.

The soil-sand-diet supplement (Sl-Sd-D), cow manure- soil-sand-diet supplement (C-Sl-Sd-D), and cow manure-sand- diet supplement (C-Sd-D) combinations all contained gravid females. A few females, when dissected, contained eggs that were full sized (Fig. 24). However, most of the females in these substrate combinations, when dissected, contained eggs in early (Fig. 25) or intermediate stages of development.

All substrate combinations containing milorganite pro- duced excessive mold growth. A 1% solution of Clorox^(© sprayed onto these mixtures as a mold inhibitor resulted in the gener­ ation of a strong ammonia-like odor. No beetles survived in these combinations.

DISCUSSION

Most ovipositional substrates evaluated did not provide adequate stimulus. Other scarabs have oviposited readily under laboratory conditions. The simple laboratory procedures outlined by Ritcher (1966) and Cartwright (1974) for inducing oviposition by some species of Aphodius and Ataenius were not applicable to A. spretulus. However, limited oviposition can be expected when inseminated females are placed on turfgrass in a manner described by Girault (1914) and Davis (1915), using P. annua or Aqrostis species as hostplants. Fig. 2k. Ovaries of A. spretulus with eggs in late stage of development.

Fig. 2 5. Ovaries of A. spretulus showing ovarioles with eggs in early stage of development. 100

REARING AND DEVELOPMENTAL STUDIES

METHODS AND MATERIALS

Field-collected A. spretulus adults, eggs, and larvae were subjected to several rearing regimes to establish a rearing method for developmental studies. Rearing conditions were maintained at 23+2 °c, 68+12% R.H., and variable photoperiod.

Illumination was necessary only for daily observation of insect development.

Field-collected eggs were placed on moist filter paper in petri dishes and incubated until larvae eclosed or signs of nonviability appeared.

Field-collected larvae and larvae from field-collected eggs were subject to 4 different rearing methods. Each method was tested in 5-10 replicates using 3-12 larvae/replicate.

One group was individually reared in 5 cm petri dishes filled with moist mixtures of soil and chopped P. annua thatch from

TPCC, plus germinating P. annua seed. A second group was placed in experimental observation rearing units (Fig. 26) filled with moist sandy loam topped with strips of P. annua sod taken from TPCC fairway 14. A third group was held in petri dishes and fed only the roots of P. annua which had been hydroponically grown (Fig. 27) in modified Hoagland's solution (Hoagland &

Arnon 1950; Christians 19 77) (Appendix 30). The fourth group was held in petri dishes containing 1-3 g of a modified, agar-base insect diet (Appendix 31). 101

# 1 1 1

Fig. 26. Observation rearing unit (a) with 10 transparent rearing troughs (b) for viewing developing larvae of A. spretulus.

>***5

Fig. 2?. Hydroponic cultures of Poa annua L. used as a source of root-diet for laboratory rearing larvae of A. spretulus. 102

Larvae developed to adults only in petri dishes filled with moist mixtures of soil, thatch and germinating P. annua seed. Therefore, this method of rearing larvae was used in a study of A. spretulus developmental rates and survivorship in which each of 9 replicates was started with a field- collected egg cluster.

Length of the pupal period was defined by placing pupae individually into small depressions in a moist peat moss-soil mixture in 9 cm petri dishes. Newly-eclosed adults were removed from pupation dishes daily and segregated by sex into containers of moist soil.

A life table was assembled for one laboratory-reared generation of A. spretulus using information generated during this study plus supplemental data from life stage sampling and overwintering studies.

RESULTS

Field-collected eggs hatched in 1 to 12 days with a surviv­ orship of 89%. First instars molted in 5 to 10 days (mean* 8 days) with 58% reaching the third instar. Third instars became prepupae in 10 to 19 days (mean * 14 days) with 58% survivor­ ship. Prepupae molted to pupae in 2 to 4 days (mean * 3 days) with a survivorship of 94%. The pupal stage lasted 7 to 10 days

(mean* 8 days) with a stadium survivorship of 100%.

Survivorship from egg to adult ranged from 8% to 25%

(mean * 17%) . Most mortality occurred among small larvae. The time from first instar to adult eclosion ranged from 28 to 55 days (mean * 41 days). If the maximum incubation period 103 observed for field-collected eggs, 12 days, approximates a

full incubation term, then the time required for A. spretulus

development from oviposition to adult eclosion would be 40 to

67 days (mean =* 53 days) at 23+2 °C and 68+12% R.H..

A separate study on the duration of adult sclerotization was conducted with 50 teneral adults from field-collected pupae. Adults became sclerotized and black in 4 to 6 days after eclosion. However, careful examination by mechancial probing of integument and inspection of spermathecae and aedeagi, revealed that complete sclerotization required at least 6 days.

These data were not sufficient for construction of a life table describing one complete generation of A. spretulus.

Data pertaining to time span and survivorship from the point of adult eclosion (prereproductive adult interval) to time of first oviposition (reproductive adult interval) had to be supplied from field studies.

The time required for an adult female to become sclerotized, copulate and become gravid was estimated at 20 days at an average temperature of 26.7+1 °C. This information was based on the time from appearance of first generation adults in sampling sites to appearance of second generation eggs

(Appendices 3-6, Figs. 9 and 10) and the average soil temper­ ature at 5 cm from June 20 to July 20, 1977 and 1978. 104

Survivorship of adult females from one age interval to

the next was not calculated using life stage sampling data

because of difficulty encountered in following movement and

distribution of individuals in fairways. Father, overwintering

survivorship was used since adults were least active and most

abundant in overwintering sites from September, 1977 through

March, 19 78. The lower limit of the estimated range (90-96%)

of overwintering survivorship was chosen for life table

assembly to allow for mortality in fairways and in transit.

A femalesmale ratio of 8:7 was used in calculating the

proportion of reproductive females and expected daughters

for the life table. This decision was based on statistical

analysis of data from life stage sampling indicating a sex

ratio slightly greater than 1:1 in favor of females.

The pooled life table (Table 21) indicates that larvae were

subject to highest mortality under laboratory conditions.

Larval mortality was respoinsible for a 72% reduction in gener­

ation size. Symptoms of moribund larvae suggested major mortality factors were desiccation, starvation and/or disease.

DISCUSSION

The small size of all life stages precluded use of rearing

containers that permitted direct observation of individuals.

The major difficulty in rearing A. spretulus in the laboratory was hesitancy of adults to mate and oviposit. Beetles rarely exhibited oviposition behavior in surroundings which differed from natural turfgrass. 105

Table 21. Life table for a laboratory-reared generation of A. spretu! 68±12^ R.H;, starting with field-collected eggs and using data from 1: overwintering studies to estimate time span and survivorship from the the reproductive adult interval.

X T. lx mx 1 xmx i

Age--Specific Survivorship Age Mean No. of Reproduct.i vc Interval Days Required Survival Rate Survival Rate Daughters Expectation for Interval No. Alive Within x from Egg

Egg 12 1 0 * 1 0 .8? 0.89 0 0

Instar 1 8 93 0.57 0.51 0 0

Tnstar 2. 8 53 0.58 0.30 0 0 o CO Instar 3 l'+ 31 wt 0.17 0 0

Prepupa 3 18 o . 9h 0.16 0 0

Pupa 8 17 1 .00 0.16 0 0

Prereprodiic ti ve Adult, 20 17 0.90 O.l'l 0 0

Reproductive Adult 0 15 0.53 0.08 0 0

Reproductive Females 0 8 - 13 1 . 0*1-

Generation Survivals 8% Mean Generation Times 73 flays Mortali tyi 92%, Net Replficpment Rate (RQ )s 1.0^1 105

table for a laboratory-reared generation of A. spretulus held at 23±2 °C, rting with field-collected eggs and using data from life stage samples and udies to estimate time span and survivorship from the prereproductive to adult interval.

lx mx l.xmx dxF dx lOOrx

Age -Specific Survivorship of Expected Reproductive Probable Number Percent i red Survival Rate Survival. Rate Daughters Expectation Mortality Dyi ng Mortali ty val No. Alive Within x from Egg Factors

10*1 0.89 0.89 0 0 Desiccation 11 11 or disease 93 0.57 0.51 0 0 Des.iccatj on, *10 '13 starve tion, or disease 53 0.58 0.30 0 0 • » It it 22 *12

ir 11 *1 11 0 . 50 0.17 0 0 13 '12

18 0.9't 0.16 0 0 Di sease 1 5

17 1 .00 0.16 0 0 - 0 0

17 0.90 0 .1*1 0 0 Di nease, 2 12 preda tion, or cold

15 0.53 0 . 0 8 0 0 Sex (99itfrf 7 53 - 8:7)

8 - - 13 1 . 0*1 Disease, 8 100 predatj on, or old age

It 8fc Mean Generation Time: 7'J days 92% Net Replacement Rate (Rq )i l.

Perhaps greater success in rearing might be achieved by

introducing overwintered adults to large screened-in plots of

P. annua or Aarostis spp. established out-of-doors or in green­

houses. Screen enclosures should be large, since adult flight

activity may be necessary for one or both sexes as a precursor

to mating and/or oviposition. Fresh turf plots may be needed

for each new generation, depending on population density. Con­ tinuous rearing may -be possible, based on observations of

Frost (1966) and Woodruff (1973) that adults have been

collected at light every month of the year in Florida.

Age interval survivorship given in the life table for the laboratory-reared generation of A. spretulus (Table 21) reflected the insect's response to artificial conditions and was not necessarily similar to that of the sampled population(s).

Desiccation, starvation and disease were the only mortality factors considered in the laboratory. However, the various species of predacious and parasitic organisms and their effects on each age interval of wild populations of A. spretulus were entirely undetermined, with the qualified exception of milky disease bacillus (species unknown).

The actual net replacement rate, Rq , for A. spretulus may be greater than that calculated for the laboratory-reared generation. If not, little or no population increase would occur from year to year. 107

The mean generation time for reared insects, 73 days, correspond well (+5 days) to first generation periods esti­ mated from life stage sampling data collected at TPCC, LCC, and KCC during 1977 (Appendices 3-5). First generation periods were determined as the number of days between appear­ ances of first and second generation eggs in samples. V. GENERAL DISCUSSION

The present study has generated the following biological information pertinent to control of A. spretulus on golf courses.

Eggs are laid during most of the growing season in Ohio due to multiple ovipositions per female and the presence of two generations. Therefore, timing of insecticide application is crucial to effective control. The phenological and thermal unit accumulation methods of timing control measures are the most accurate and will yield the best results, as demonstrated by Niemczyk and Wegner (1979).

Natural enemies alone cannot be considered adequate in a management strategy for A. spretulus on golf courses.

Due to the small size of the larvae, damaged to turf- grass becomes apparent only where infestation density reaches 2 100-600/929 cm and larvae are nearly full size. Therefore, frequent, careful checks should be made for presence of larvae in turfgrass, during late May and early June, at golf courses with no previous record of A. spretulus infestation.

Damage from the second generation can be more severe than that caused by the first if all greens, tees, and fairways

108 109 have not been treated with an effective insecticide during spring oviposition.

Infestation density of A. spretulus in turfgrass the year after it was thoroughly treated may be equal to that of the previous year. This is due to invasion by overwintered insects which developed in untreated turfgrass or nearby areas in the previous year. VI. SUMMARY

A. spretulus the black turfgrass ataenius, is an important

pest of golf course greens, tees and fairways in Hamilton and

Clermont counties, Ohio. Eggs average 0.75 x 0.52 mm and are

deposited in clusters of 11-12 in the thatch and thatch-soil

interface. Head capsule widths of larvae average 0.5 mm for

first instars, 0.83 mm for second instars and 1.3 mm for third

instars. Mature larvae average 8.5 mm long and have 40-45

irregularly placed, hamate setae on their rasters. The root-

feeding larvae have caused wilt or death of host turfgrass,

P. annua and Aarostis species, in infestations of 100-600/929 2 cm . Pupae average 4.7 x 2.5 mm and occupy cells excavated 1-8

cm deep in the soil. Adults average 4.9 x 2.2 mm and are shiny black. Adults are weak fliers but are common in flight over

fairways on warm, sunny afternoons in early spring and fall

and around lights on summer evenings.

Evidence for two generations of A. spretulus per year was strong at the 3 courses sampled. The appearance of teneral

adults and increased numbers of mature adults collected from

life stage samples and light traps in late June and early July,

1977 and 1978, indicated the completion of the first generation.

110 Ill

An abrupt increase in the number of eggs found in samples

around mid-July, following a gradual decline, during June and

early July, indicated the beginning of a second generation.

Perennial differences in A. spretulus population size on

golf courses and seasonal fluctuations in infestation density

between first and second generations were controlled by one or

more factors: (1) dispersal due to diminished food in the

preferred habitat, (2) abundance of predators and parasites,

and (3) frequency and extent of insecticide application on

(turfgrass) playing surfaces.

Between-course and within-course comparisons of life

stage sampling data indicated that populations at the 3 courses

were distinct statistically, though not necessarily

biologically.

The dispersion pattern of A. spretulus infestations in the

sampling sites best conformed to a highly contagious or

logarithmic distribution. This may have resulted from the dis­

criminating behavior of ovipositing females in seeking the

favored microclimate and the laying of eggs in clusters.

Sex ratios of adults and pupae sampled during 1977 and

1978 exhibited significant variation (P>.05) from 1:1 in only

17% of monthly records. Although statistical significance was infrequent, females tended to outnumber males slightly.

The inseminated fraction of females taken in life stage

samples, light traps, and sticky traps during 1977-1978 was 112

significantly greater (P> .05) than the virgin fraction in

every month except July. Most of the second generation females

collected during August-October were inseminated before migrat­

ing to suitable overwintering sites. Overwintered adults from

the previous year's second generation (most of which were

inseminated) were the source of all the females collected in traps and samples from April to June. July samples and trap

captures contained significantly fewer inseminated females than

other months because many of the adults present at that time were newly-eclosed, especially early in July, and had not yet

copulated.

Adults of A. spretulus overwintered in greatest abundance beneath duff and ground cover, in 1 to 5 cm of soil at the tree line of wooded areas located near host turfgrass. Up to 264 2 adults/929 cm were found in these locations from September through March. Overwintering survivorship was estimated at

90 to 96%. There was no significant difference (P>.05) between survivorship of females and males.

Populations of A. spretulus are most vulnerable to control by insecticide in early to mid-May when overwintered, gravid females seek oviposition sites, as suggested by Niemczyk and

Wegner (1979). This time corresponds to first full bloom of

Vanhoutte spirea, horse chestnut, and to first bloom of black locust (Niemczyk & Wegner, 1979). The phenological method of predicting the appropriate time to apply insecticide is much more accurate than calendar dating, year after year, because both insect and plant activity are correlated through heat unit 113

accumulations. The flight threshold of A. spretulus was

estimated to be 13 °C and was used instead of the developmental

threshold to calculate day degree accumulations for major

points in its seasonal history.

The effect of natural enemies of A. spretulus infestations

at Ohio sampling sites was insufficient to prevent extensive

damage to turfgrass during the two year sampling program.

Larvae at all 3 courses exhibited symptoms typical of infection

by a milky spore bacterium. Incidence of milky disease at

TPCC, LCC, and KCC ranged from <1% in May and June to 25-29%

in October, 1977. Milky disease infection of larvae at TPCC

in 1978 ranged from 0% in May to 1% in September.

Other parasites found in larvae were a nosematid micro-

sporidian, a neoaplectanid nematode, a cephaline eugregarine,

and an Hyphomycetes fungus.

No insects were observed feeding on A. spretulus in

turfgrass. However, 3 species of carabid beetle were found in

that habitat and could have preyed on immatures. Amara

lecontei (Csiki), Amphasia interstitialis (Say), and

Harpalus fulgens Csiki. The starling was probably the most

important vertebrate predator of A. spretulus.

The following mites were phoretic on migrating adults:

Parasitus sp., Dendrolaelaps sp., Scutacarus sp., Histiostoma

sp., Macrocheles sp., and Macrocheles insiqnitus (Berlese).

Results of laboratory rearing studies indicate that

approximately 5 3 days were required for development from 114 oviposition to adult eclosion at 2 3+2 °c (Table 2 1 ) . If, based on field observations, 20 days are added to allow for mating and egg maturation in females, the time required for a second generation would be about 73 days. However, less time may have been required for development in the field since the soil temperature 5 cm deep often approached or exceeded 30 °C during July and August, thereby accelerating the rate of development.

Beginning with oviposition on May 1, a generation period of 60-70 days would place first generation adult eclosion in late June and second generation adult eclosion in late

August and early September. This schedule conforms well to* an observed increase in the proportion of teneral and mature adults found in early July and late August.

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131 132

APPENDIX 1

Analysis of soil from 4x3 m sites at each golf course sampled for overwintering survival of A. spretulus adults.

. . Percentile Composition Site pH Organic Matter Sand Silt Clay

TPCC 7.3 1 72 14 13 LCC 6.9 3 43 26 28 KCC 5.8 4 50 20 26 APPENDIX 2

Standard fairway treatments at 3 golf courses where A. spretulus life stage sampling was conducted from 1977 through 1978.

Category Description Locati on TPCC LCC.. KCC Mowing + + + iH » Coring + + + o, Irrigation + + + <8> Acti-Dione TFG (cycloheximide) + _ + Daconii!® (chlorothalonil tetra- + + - to a> chloroisophthalonitrile) T> •H Chloroneb (1,4-dichlor0-2,5- + O •H dime thoxy-benzene ) Lesco-4(zinc ion, manganese + + - ethylene bisdithiocarbamate)

FMA (phenyl mercuric acetate) + + -

Thiram (tetramethylthiuram + - - disulfide) Betasan®(S-(o,o-diisopropyl — — phosphorodithi oate) of N-(2-mercaptoethyl) benzene- sulfonamide) § DacthaiPw-75 (dimethyl tetra- + + chloroterephthalate) MCPP (2-[(4-chloro-o-tolyl)oxy] - + propionic acid * Trimec (diethylamine salts of + 2,4-dichlorophenoxyacetic acid 2-(2-methyl-4-chlorophenoxy) propionic acid dicamba (3,6- dichloro-o-anisic acid))______Milorganite® - + Scott HD Fairway Fertilizer® + + | (10-1-3) £ Phosphorus (0-46-0) + +» Ferrous sulfate + a Sulfate of potash (0-0-50) + Ureaform (38-O-O) + APPENDIX 3

Number of each life stage of A. spretulus found per sampling occasion at TPCC during 1977.

Numbers/Age Class/Sampling S i t e ^ Oats Adults 1 i Instar 3 Prenunae Punas l4 17 14 17 17 it 17 14 17 14 17 14 17 4/i4 2 3 0 0 0 0 0 0 0 0 0 0 0 0 19 2 0 0 0 0 0 0 0 0 0 0 0 0 0 22 2 1 0 0 0 0 0 0 0 0 0 0 0 0 26 2 0 0 0 0 0 0 0 0 0 0 0 0 0 29 4 2 0 0 0 0 0 0 0 0 0 0 0 0 5/ 3 3 1 0 12 0 0 0 0 0 0 0 0 0 0 6 2 3 16 0 0 0 0 0 0 0 0 0 0 0 10 2 1 45 17 0 8 0 0 0 0 0 0 0 0 13 1 2 0 11 0 0 0 0 0 0 0 0 0 0 17 1 1 1 21 1 7 0 0 0 0 0 0 0 0 20 1 0 1 0 2 0 1 3 0 0 0 0 0 0 2>* 0 0 20 8 4 0 15 17 0 0 0 0 0 0 27 0 0 1 20 3 3 35 32 18 2 0 0 0 0 6/ 1 0 0 0 9 0 4 0 5 12 43 0 0 0 0 3 0 1 10 8 5 0 4 2 106 45 0 0 0 0 7 0 0 0 5 0 6 2 9 33 67 17 10 0 0 10 0 0 0 4 3 5 12 48 107 65 32 10 0 0 14 0 0 0 0 0 1 18 16 34 49 61 20 20 5 17 0 0 0 12 1 5 17 32 48 34 24 19 37 21 21 0 0 0 12 3 18 5 25 23 41 8 34 55 11 24 17 3 0 22 0 4 1 5 19 80 23 19 33 61 28 23 4 0 0 0 0 8 10 45 32 25 16 54 35

7/ 1 24 15 0 0 6 4 2 12 16 77 9 14 40 32 6 23 34 0 0 2 0 2 0 8 17 5 6 15 ll 134 3 39 33 0 0 0 0 0 10 4 25 4 2 15 12 13 14 33 47 0 0 0 9 1 21 44 4 3 17 14 15 18 19 24 0 1 0 9 10 9 34 3 2 8 5 18 17 28 0 12 5 1 14 7 5 25 0 2 2 17 20 9 28 12 20 7 2 16 2 14 21 0 2 2 5 25 4 5 19 0 0 3 21 13 44 3 2 2 0 5 27 6 29 0 0 6 0 15 13 11 26 2 1 2 4 8 22 12 23 2 2 16 0 8/ 19 5 39 1 2 7 4 9 0 8 1 4 3 17 66 29 4 7 0 0 13 1 2 0 11 0 0 0 0 0 0 0 0 0 0 17 1 1 1 21 1 7 0 0 0 0 0 0 0 0 20 1 0 1 0 2 0 l 3 0 0 0 0 0 0 2b 0 0 20 8 b 0 15 17 0 0 0 0 0 0 27 0 0 1 20 3 3 35 32 18 2 0 0 0 0 6/ 1 0 0 0 9 0 b 0 5 12 b3 0 0 0 0 3 0 1 10 8 5 0 b 2 106 b5 0 0 0 0 7 0 0 0 5 0 6 2 9 33 67 17 10 0 0 10 0 0 0 b 3 5 12 b8 107 65 32 10 0 0 lb 0 0 0 0 0 1 18 16 3* b9 61 20 20 5 17 0 0 0 12 1 5 17 32 b8 3b 2b 19 37 21 21 0 0 0 12 3 18 5 25 23 bl 8 3b 55 11 2b 17 3 0 22 0 b 1 5 19 80 23 19 33 61 28 23 b 0 0 0 0 8 10 <►5 32 25 16 5b 35

7/ 1 2b 15 0 0 6 b 2 12 16 77 9 lb bo 32 6 23 3b 0 0 2 0 2 0 8 17 5 6 15 11 3 39 33 0 0 0 0 0 10 b 25 b 2 15 12 134 13 lb 33 b7 0 0 0 9 1 21 bb b 3 17 lb 15 18 19 2b 0 1 0 9 10 9 3b 3 2 8 5 18 17 28 0 12 5 1 lb 7 5 25 o 2 2 17 20 9 28 12 20 7- 2 16 2 lb 21 o 2 2 5 25 b 5 19 0 0 3 21 13 bb 3 2 2 0 5 27 6 29 0 0 6 0 15 13 11 26 2 1 2 b 8/ 1 8 22 12 23 2 2 19 5 39 16 1 0 2 7 3-bS/ b 9 0 8 1 b 3 17 66 29 b 7 0 0 6 7 10 0 0 0 13 1 16 36 bb 5 1 7 0 10-11^/ 7 19 2b 36 2 b 9 bl 21 5* 6 2 7 2

15 10 2 22 29 1 5 12 28 7 61 1 10 12 2 17 5 b 0 11 0 0 0 12 15 37 b 8 3 10 22 8 8 0 0 0 0 b 13 7 57 1 2 3 3 2b 20 b 11 0 0 6 3 9 15 65 2 6 18 lb 30 7 11 0 0 0 b 0 b lb 58 2 b 1 b

9/ 2 5 16 0 11 2 0 1 10 18 60 2 3 5 16 7 6 15 11 0 0 0 0 b 12 33 1 10 6 15 9 6 10 0 0 0 0 2 0 10 29 6 2 6 10 12 b 17 0 0 0 0 0 3 2 13 0 b b 17 lb 3 20 0 0 0 5 0 6 10 27 7 b 3 20 19 b 7 0 0 0 0 0 1 7 8 2 3 b 7 21 3 12 0 0 0 0 0 1 2 22 2 5 3 12 26 2 6 0 0 0 0 0 1 3 12 0 0 2 6 10/ 3 1 5 0 0 0 0 0 1 b lb 3 0 1 5 13 2 2 0 0 0 0 0 1 3 1 0 1 2 2 18 2 10 0 0 0 0 5 1 6 11 2 2 2 10

a. Sampling sites designated by fairway number. b. Two-day period considered as one sampling occasion. APPENDIX 4

Number of each life stage of A. spretulus found per sampling occasion at LOG during 1977*

Numbara/Aga Claas/Sampling 51ta*/ Data A d u l t s Inatar 1 Ina t a r 2 Inatar 1 bunaa 11 16 11 16 11 16 11 16 11 16 11 16 11 16

9 /15 2 1 0 0 0 0 0 0 0 0 0 0 0 0 16 O 1 0 0 0 0 0 0 0 0 0 0 0 0 20 1 1 0 0 0 0 0 0 0 0 0 0 0 0 25 9 3 0 0 0 0 0 0 0 0 0 0 0 0 27 2 1 00000000000 0 5/ 2 2 0 10 20 0 0 0 0 0 0 0 0 0 0 5 3 3 10 21 0 0 0 0 0 0 0 0 0 0 9 0 1 15 52 0 0 0 0 0 0 0 0 0 0 11 0 2 0 20 0 0 0 0 0 0 0 0 0 0 19 0 1 22 0 9 0 0 0 0 0 0 0 0 0 IS 3 0 32 13 5 12 0 0 0 0 0 0 0 0 23 0 1 97 9 31 0 6 0 0 0 0 0 0 0 25 0 i_ 21 27 6 15 33 9 0 0 0 0 0 0 31 1 0 10 0 7 9 22 29 15 8 0 0 0 0 6 / 3 1 0 12 98 18 17 32 99 85 a 3 0 0 0 6 1 I 0 0 12 10 12 26 93 50 0 0 0 0 e 1 0 23 0 17 a 91 26 99 18 1 0 0 0 13 0 0 10 0 17 6 90 32 86 50 7 0 0 0 16 0 0 8 3 0 I 37 36 111 53 7 2 0 0 21 0 0 11 8 9 0 29 10 76 37 27 19 3 9 23 0 0 0 0 0 3 3 0 60 33 52 30 31 11 27 0 0 0 0 0 0 5 2 79 61 31 13 26 23 30 2 0 0 0 2 0 3 3 58 30 91 5 30 J2/ 7/ 5 9 10 0 0 5 0 10 1 16 93 7 9 93 16 7 15 17 0 5 0 2 11 3 17 39 19 9 92 27 12 38 7 0 7 0 9 1 3 12 17 9 0 12 22 15 92 12 0 12 0 0 9 0 9 11 0 3 9 6 19 6 13 0 0 0 9 0 0 9 12 0 9 7 15 21 8 37 0 0 0 0 0 3 3 21 0 3 1 25 26 13 17 0 0 0 2 2 13 1 13 9 1 1 2 23 19 9 0 11 0 0 8 1 0 a 1 0 3 1 8/ 2 11 12 0 13 0 1 10 33 l 32 0 2 0 3 5 6 11 0 5 5 5 0 9 0 29 1 0 i 0 9 10 7 9 9 7 9 9 6 10 23 0 2 2 17 11 1 13 0 12 2 19 9 12 28 15 0 2 0 9 16 6 2 12 37 0 5 9 8 13 31 1 3 2 9 19 3 0 0 27 0 11 2 20 9 23 3 3 I 2 23 * 3 0 0 0 9 1 23 6 23 2 0 5 12 26 9 6 0 0 6 9 3 7 12 10 2 5 1 21 29 1 6 9 0 0 0 0 6 11 16 0 2 2 2 31 9 1 0 0 1 I 6 0 12 11 8 2 2 9

9/ 6 29 2 0 0 0 0 1 1 19 19 a 1 19 3 3 2 7 0 0 0 0 0 2 8 31 2 1 3 6 13 £ 1 0 0 0 0 1 0 13 0 1 0 0 3 16 2 2 0 0 0 0 0 0 11 9 0 3 1 13 20 0 5 0 0 0 0 0 0 12 9 3 2 10 8 23 12 9 0 0 0 0 3 0 7 9 9 2 11 2 26 3 9 0 0 0 0 0 0 9 5 2 1 12 0 1 0 / 5 9 6 0 0 0 0 0 0 6 6 0 0 3 8 11 1 9 0 0 3 0 0 0 3 1 1 1 0 7 20 6 5 0 0 0 0 0 0 1 9 1 7 3

a. Sampling aitaa daaignatad by fairway numbar. b. Missing datum. APPENDIX 5

Number of each life stage of A. spretulus found per sampling occasion at KCC during 1977.

Huabars/Aga'C3ag«/Samallnit Slt>^ gtia lg Inatar ^ Inatar | IjMtar j f^ePUBBg______S fUBM4

4 /14 0 0 0 3 0 0 0 0 0 0 0 3 0 0 13 0 1 0 0 0 3 0 3 3 3 3 3 0 0 20 0 0 0 0 0 0 0 0 0 0 3 0 0 0 26 1 0 0 0 0 0 3 0 0 0 3 0 0 0 29 2 2 0 0 0 0 0 0 0 0 3 0 0 0 5/ 2 0 0 0 0 0 0 3 0 0 0 0 0 0 0 * 0 1 0 0 0 0 0 0 3 0 0 0 0 0 3 0 0 10 0 0 0 0 0 0 0 3 0 0 0 12 0 1 0 12 0 0 0 0 0 0 3 0 0 3 I? 0 0 0 3 0 3 3 0 3 3 0 3 0 0 1? 3 0 0 0 0 0 0 0 0 0 0 0 0 0 23 1 3 0 0 0 0 0 0 0 0 0 0 0 0 26 0 0 a 0 0 0 0 0 0 0 3 0 0 0 31 0 0 a 0 0 0 2 2 9 1 3 3 0 0 6/ 2 0 0 a 0 0 1 3 12 2 9 0 0 0 0 5 0 0 0 0 0 0 4 3 7 3 0 0 0 1 0 1 a 0 0 1 13 0 35 4 5 0 0 0 13 2 3 0 0 16 7 9 4 53 9 10 0 0 0 15 0 3 0 0 4 « 4 14 47 14 3 0 0 a 30 0 0 0 0 0 2 1 3 29 24 6 2 4 0 22.23*/ 3 0 0 10 0 0 9 12 37 73 14 2 17 i 27 1 1 0 0 0 0 0 2 13 44 6 £ 16 10 t 29 ! 1 0 0 0 3 0 0 3 36 6 11 13

7/ 5 13 a 0 0 0 0 3 4 12 36 3 4 16 5 7 17 0 0 0 0 0 0 1 7 5 1 0 5 7 12 5 3 0 0 0 0 0 1 6 19 0 4 V7 30 lit 5 2 0 0 0 0 0 0 1 24 6 2 10 5 19 5 12 12 24 7 0 0 l 0 18 0 6 0 13 22 7 14 to 0 0 0 9 0 3 13 0 7 0 10 26 7 11 10 11 0 0 3 £ 1 9 0 1 0 6 29 4 13 0 3 0 0 3 1 14 5 0 2 0 12 8/ 1 1 16 0 0 1 7 13 13 21 13 0 0 1 6 4 2 17 0 0 0 1 10 7 43 20 0 0 1 3 10 3 16 22 0 4 2 14 35 37 41 4 3 7 11 12 2 20 0 0 3 1 9 41 55 76 4 10 4 17 15' 0 14 0 32 0 0 4 3 53 43 3 2 5 10 13 1 19 0 36 0 0 V 7 49 50 13 10 16 22 23 3 4 0 11 2 0 11 7 20 44 13 10 10 15 25 13 13 0 0 0 3 3 3 21 23 11 5 17 30 30 15 22 11 0 0 0 3 0 30 32 7 6 14 24 9/ 1 12 19 0 0 0 0 0 0 4 10 4 5 11 19 £ 11 30 0 0 0 0 i 4 26 8 1 0 5 21 3 ■ M 15 0 0 0 0 0 3 21 25 4 1 5 3 12 13 12 0 0 0 0 0 0 > 0 2 0 2 3 15 17 13 0 3 0 0 0 0 2 2 4 3 4 ? 19 9 22 0 3 a 0 1 3 n 3 1 0 6 1 22 5 10 0 0 0 0 0 0 9 t 4 1 4 3 30 i 1 0 0 0 0 0 3 2 7 2 1 0 0 10/ 7 4 9 3 3 0 0 3 0 4 3 3 0 7 3 1 I 1 0 0 0 0 0 0 0 I 0 1 7 3 18 3 5 a 0 0 0 0 3 2 2 1 1 3 1

a. Sampling sitas designated by fairway numbar.

b. Two-day pariod eonsidarad as ona sampling occasion. APPENDIX 6

Number of each life stage of A. spretulus found per sampling occasion at TPCC during 1978.

a/ Numbers/Aae Class/Sampling Sit&k/ Date --- “- — ------. Adults______E g g s ______Instar 1 InsSatyS-----lagtar. 3 - Preoupae Punae i4b 14a i 4b iEa Tub i4a i 4b I4a i 4b i4a i 4b 14a i 4b 4/ 5- 6 2 0 0 0 0 0 0 0 0 0 0 0 0 0 12-13 6 2 0 0 0 0 0 0 0 0 0 0 0 0 19-20 14 1 0 0 0 0 0 0 0 0 0 0 0 0 26-27 7 6 0 0 0 0 0 0 0 0 0 0 0 0 5/ 2- 3 3 5 0 0 0 0 0 0 0 0 0 0 0 0 10-11 16 3 0 0 0 0 0 0 0 0 0 0 0 0 17-18 25 5 0 0 0 0 0 0 0 0 0 0 0 0 24 2/ 10 72 - 0 - 0 - 0 - 0 - 0 - 5/31-6/1 4 3 19 9 10 4 0 0 0 0 0 0 0 0 6/ 7 - 8 2 10 25 0 25 10 75 121 0 17 0 0 0 0 14-15 0 1 24 0 13 0 64 25 67 199 0 0 0 0 21-22 1 l 11 0 2 0 24 4 159 281 4 14 0 0 28-29 1 3 0 0 1 0 9 9 99 94 43 76 24 103 7/ 5- 6 8 30 22 20 0 0 4 0 35 26 12 51 52 1*5 12-13 46 75 0 0 0 0 9 1 18 10 4 6 22 70 19-20 38 68 19 22 8 0 2 2 15 3 0 7 9 22 26-27 38 82 57 12 6 0 2 5 19 15 1 1 13 3

8/ 2- 3 60 50 41 22 7 9 8 8 14 11 6 1 0 2 9-10 21 10 30 49 12 1 52 3 66 8 1 8 3 16-17 15 16 0 0 2 5 15 8 48 13 0 1 2 6 23-24 21 18 0 6 4 0 31 3 68 35 3 1 17 6 30-31 23 7 0 12 2 3 8 10 ' 64 22 8 1 25 8 9/ 6- 7 32 23 0 0 0 0 0 2 31 29 10 1 30 12 13-14 28 21 0 21 0 1 2 3 12 12 2 3 9 15 20-21 40 34 0 0 0 1 2 0 12 7 1 3 3 7 27-28 25 19 0 0 4 0 1 0 7 14 0 2 2 5 10/ 4- 5 25 14 0 0 0 0 0 0 0 3 0 0 0 5 11-12 10 11 0 0 0 0 0 0 4 2 0 0 1 0 18-19 17 11 0 0 0 0 0 0 3 4 0 0 0 2 25 10 8 0 0 0 0 0 0 2 3 0 0 0 3

a. Two-day periods considered as single sampling occasions. b. Sampling sites designated by fairway number? A and B * opposite ends of fairway 14. From 4/5 to 5/24, area 14a = 14 and area 14B * 17. c. Missing data from fairway 17 due to inadvertent treatment withinsecticide. 138

APPENDIX 7

Sex ratio of A. spretulus adults collected from sampling sites at TPCC in 1977.

Males Females Deviation from Month No. Sexed No. Percent No. Percent 1«1 Sex P.atio (P< .05)

APR 18 7 38.89 11 61.11 .3472 n.s. MAY 17 7 41.18 10 58.82 .4654 n.s. JUN 43 20 46.51 23 53.49 .6456 n.s. JUL 36l 165 45.71 196 54.29 .1032 n.s. AUG 159 79 49.69 80 50.31 .9362 n.s. SEP 134 70 52.24 64 47.76 .6030 n.s.

OCT 22 -11 50.00 JL1 50.00 1.0000 n.s. Total 754 3 59 47.61 395 52.39 1 3 9

APPENDIX 8

Sex ratio of A. spretulus adults collected from sampling sites at LCC in 1977*

Deviation from Males Females Month No. Sexed 1il Sex Ratio No. Percent No. Percent (P< .05)

APR 16 5 31.23 11 68.75 .4532 n.s. MAY 17 0 0.00 17 100.00 .0000 « JUN 6 2 33.33 4 66.67 .4122 n.s. JU1 222 103 46.40 119 53.60 .2846 n.s. AUG 114 46 40.35 68 59.65 .0394 # SEP 66 30 45.45 36 54.55 .4592 n.s. OCT 26 -11 51-37 JL2 46.4”? .7040 n.s. Total 469 201 42.95 268 57.05 140

APPENDIX 9

Sex ratio of A. spretulus adults collected from sampling sites at KCC in 1977*

Deviation from Males Females Month No. Sexed No. Percent No. Percent 1:1 Sex Ratio (P< .05)

APR 6 3 50.00 3 50.00 1.0000 n.s. MAY 6 2 33.33 4 66.67 .4122 n.s. JUN 9 3 33.33 6 66.67 .3174 n.s. JUL 116 50 43.10 66 56.90 .1362 n.s. AUG 174 96 55.17 78 44.83 .1738 n.s. SEP 197 95 49.24 102 50.76 .6170 n.s. OCT 40.91 _n 59.09 .3954 n.s. Total 530 258 48.68 272 51.32 141

APPENDIX 10

Sex ratio of A. spretulus pupae collected from sampling sites at TPCC in 1977.

Month No. Sexed Males Females Deviation from No. Percent No. Percent 1 «1 Sex Ratio (P< .05)

JUN 302 127 42.05 175 57.95 .0058 * JUL 198 65 32.83 133 67.17 .0000 * AUG 92 48 52.17 44 47.83 n.s. SEP 97 46 47.42 51 52.58 .6100 n.s. OCT _ 2 33.33 6 66,67 .3174 n.s. Total 698 289 41.40 409 58.60 142

APPENDIX 11

Sex ratio of A. spretulus pupae collected from sampling sites at LCC in 1977-

Deviation from Males Females 1 il Sex Ratio Month No. Sexed No. Percent No. Percent (P< .05)

JUN 135 69 51.11 66 48.89 .7948 n.s. JUL 226 91 40.2? 135 59.73 .0034 * AUG 84 44 52.38 40 47.62 .6600 n.s. SEP 85 44 51.76 41 48.24 .7414 n.s. OCT _g8 JL5 531.57 _n 46.43 .7040 n.s. Total 558 263 47.13 • 295 52.87 143

APPENDIX 12

Sex ratio of A. spretulus pupae collected from sampling sites at KCC in 1977. Deviation from Males Females Month No. Sexed 1 tl Sex Ratio No. Percent No. Percent (P< .05)

JUN 72 35 48.61 37 51-39 .8104 n.s. JUL 125 58 46.40 6? 53.60 .4238 n.s. AUG 199 76 38.19 123 61.81 .0008 * SEP 94 43 45.74 51 54.26 .4066 n.s.

OCT _LZ 6 3.5-2? 11 64.71 .2262 n.s. Total 50? 218 43.00 289 57.00 APPENDIX 13

Sex ratio of A. spretulus adults collected from sampling sites at TPCC in 1978.

Deviation from Male Female 111 Sex Ratio Month No. Sexed Percent No. Percent (P <. 05)No.

APR 38 18 47.37 20 52.63 .7490 n. s. MAY 71 28 39.44 43 60.56 .0750 n.s. JUN 22 8 36.36 14 63.64 .2006 n. s.

JUL 38 5 208 54.03 177 45.97 .0000 * AUG 2*4-0 154 64.17 86 35.83 .0000 * SEP 221 99 44.80 122 55.20 .1212 n.s. OCT 103 _56 54.37 _42 45.63 .3844 n. s. Total 1080 571 52.87 509 47.13 145

APPENDIX 1^

Sex ratio of A. srretulus pupae collected from sampling sites at TPCC in 1978.

Deviation from Males Month No. Sexed Females lil Sex Ratio No. Percent No. Percent (P <.05)

JUN 127 66 51.97 61 48.03 .6600 n.s. JUL 336 169 50.30 167 49.70 .9124 n.s. AUG 77 41 53-25 36 46.75 .5686 n.s. SEP 83 37 44.58 46 55.42 .3222 n.s. OCT _11 __2 27.27 72.73 .1310 n.s. Total 634 316 49.84 318 50.16 146

APPENDIX 15

Number of A. spretulus adults captured per day on rectan­ gular screen sticky traps at TPCC during 1977•

Date APR MAY JUN JU1 . AUG SEP OCT I'*a/17 14 17 14 17 14 17 w 17 14 L7 14 17 1 0 0 0 6 0 0 0 0 0 6 2 6 2 1 0 0 0 0 0 0 0 0 0 3 0 3 0 0 0 0 0 0 0 0 0 0 0 0 4 1 0 0 0 1 0 2 0 0 0 0 0 5 0 0 0 0 1 0 0 0 2 1 0 0 6 0 0 0 0 3 0 0 0 1 0 1 0 7 0 0 0 0 0 0 1 0 0 0 0 0 8 0 0 0 0 2 0 0 0 0 0 0 0 9 0 0 0 0 1 1 0 0 0 0 0 0 10 0 0 0 0 0 0 1 0 0 0 0 0 11 0 0 0 0 10 0 2 1 0 0 0 0 12 - 5/ - 0 0 0 0 3 0 0 0 0 0 0 0

13 6 0 • - 0 0 15 1 5 0 6 0 1 0 • lit 13 0 • 0 0 0 0 3 0 0 0 1 0 {£/ 15 3 0 - 0 0 2 0 0 0 5 1 0 0 16 4 0 0 0 0 0 4 0 0 0 0 0 0 0 17 0 0 1 0 0 0 2 0 0 0 4 0 0 0 18 0 0 0 0 0 0 3 1 0 0 5 0 0 0 19 0 0 0 0 0 0 2 0 0 0 0 0 0 0 20 0 0 0 0 0 0 1 1 0 0 0 0 0 0 21 1 0 0 0 0 0 0 0 0 0 0 0 2 0 22 0 0 1 0 0 0 0 0 1 0 0 0 0 0 23 0 0 0 0 0 0 0 0 1 1 1 0 0 0 24 0 0 0 0 0 0 2 0 0 0 1 0 0 0 25 0 0 0 0 0 0 2 0 0 0 0 0 0 0 26 0 0 0 0 0 0 0 0 0 0 1 0 1 0 27 0 0 0 0 0 0 0 0 1 0 0 0 0 2 28 0 0 0 0 0 0 9 0 1 0 0 0 0 0 29 0 0 0 0 0 0 1 0 0 0 0 0 0 0 30 0 0 0 0 0 0 3 0 0 0 4 0 0 0 31 0 0 4 0 0 0 0 0

a. Traps designated by number of adjacent fairway. b. Missing datum. c. Datum preceded by (....) indicates cumulative data for that period of time. 147

APPENDIX 16

Number of A. spretulus adults captured per day on rectan­ gular screen sticky traps at LCC during 1977.

Date APR MAY JUN JUL AUG SEP OCT H&/ 16 11 16 11 16 11 16 11 16 11 16 11 16

1 0 0 0 0 0 1 0 0 0 1 7 45 2 0 3 0 0 0 0 0 0 0 0 0 2 3 0 3 0 0 0 0 0 2 0 0 0 0 4 0 3 0 0 0 1 0 4 0 0 0 0 5 0 3 0 0 0 4 1 2 0 0 0 0 6 0 L 0 0 0 0 2 2 0 0 0 0 7 0 3 0 0 0 1 0 6 0 0 0 0 8 0 3 0 0 0 0 0 2 0 0 0 0 9 0 3 0 0 0 0 2 2 0 0 0 0 10 0 3 0 0 0 0 1 13 0 0 0 0 11 0 3 0 0 1 0 1 1 0 0 0 0 12 . 22/ - 0 3 0 0 0 1 0 6 0 0 0 0 13 15 - 0 0 0 9 2 12 0 0 0 0 14 - 9 - 0 1 0 19 3 50 0 0 0 0 15 0 2 - ■V 0 0 0 7 0 3 7 0 0 0 16 0 1 1 1 0 0 0 4 0 2 3 4 0 0 17 0 2 0 1 0 0 0 13 0 0 1 4 0 0 18 0 4 0 0 0 0 0 9 0 0 0 6 0 0 19 0 0 0 0 0 0 0 7 0 0 0 0 0 0 20 0 1 0 0 0 0 1 9 0 0 3 0 2 0 21 0 0 0 0 0 0 0 1 1 0 0 0 5 6 22 0 0 0 0 0 0 0 0 1 1 0 0 0 0 23 0 0 0 3 0 0 0 0 3 10 5 3 0 2 24 0 0 0 0 0 0 0 0 1 0 1 0 0 1 25 0 0 0 0 0 1 0 1 0 0 3 0 0 0 26 0 0 0 0 0 0 0 0 4 5 0 0 0 0 27 0 0 0 0 0 0 0 4 6 0 0 0 2 28 0 0 0 0 0 0 0 1 2 0 0 0 0 29 0 0 0 0 0 0 1 0 0 0 0 0 0 30 0 2 0 0 0 0 3 0 1 s 4 0 0 31 0 0 p 1 . 9_____2 p 9

a. Traps designated by number of adjacent fairway. b. Missing datum. c. Datum preceded by (....) indicates cumulative data for that period of time. 148

APPENDIX 17

Number of A. spretulus adults captured per day on rectan gular screen sticky traps at KCC during 1977.

Date APR______MAJ______JUN JUL AUG . SEP OCT 4 & 6 4 6 4 6 4 6 4 6 4 6 4 6 1 0 0 0 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 3 0 0 0 0 0 0 0 2 0 1 0 0 4 0 0 0 0 0 0 2 4 0 1 0 0 5 3 0 0 0 5 0 1 3 0 1 0 0 6 0 0 0 0 3 1 1 1 0 0 0 0 ? 0 0 0 0 0 0 1 1 0 0 0 0 8 0 0 0 0 0 0 4 8 0 0 0 0 9 0 0 0 0 0 0 11 2 0 1 0 0 10 0 0 0 0 0 6 5 0 0 0 0 11 o o o : 0 0 4 3 0 0 0 0 12 - ^ 5 o 0 o i 5 0 0 3 0 0 0 0 13 4i 3 o.o o o 9 2 1 7 0 0 0 0 14 4 11 0 0 0 0 0 1 4 2 0 0 0 1 15 o i o o o o 0 0 2 4 0 2 0 0 16 2 13 0 0 0 0 0 1 2 6 0 0 0 0 17 0 0 1 2 2 0 2 1 0 0 0 1 0 0 18 0 1 2 0 0 0 2 3 0 0 1 3 0 0 19 0 0 0 0 0 0 1 0 1 0 0 0 0 1 20 i i: c/ : o o 0 0 0 0 0 0 0 3 21 0 0 3 : 0 0 2 0 0 0 0 0 2 7 22 0 0 o i 0 0 0 0 0 1 0 0 0 0 23 0 0 0 0 0 0 0 0 2 11 0 3 0 0 24 0 0 0 0 0 0 2 3 0 0 0 1 0 0 25 0 0 0 0 0 0 0 0 0 0 1 1 0 0 26 0 0 0 0 0 0 0 0 0 1 0 0 0 0 27 0 0 0 10 0 0 0 0 0 0 0 2 6 28 0 0 0 * 0 0 2 2 0 1 0 0 0 0 29 0 0 0 * 0 0 1 0 o' 0 0 0 0 0 30 _0___0 0 i _0___0 0 0 0 0 0 0 0 0 31 _0___ 0 10 Z 0 z 0 0

a. Traps designated by number of adjacent fairway. b. Missing datum. c. Datum preceded by (....) indicates cumulative data for that period of time. APPENDIX 18

Maximum, minimum, and mean numbers of A. spretulus adults captured per day on rectangular screen sticky traps at TPCC, LCC, and KCC during 1977.

APR MAY JUN JUL AUG SEP OCT

Date Max*/ £/x*/ Max X Max X Max X Max X Max X Max X

1 0 0 0 0 0.27 0.06 0 0 0.29 0.06 12.10 3.97 2 0.81 0.26 0 0 0 0 0 0 0 0 0.81 0.32 3 0 0 0 0 0 0 0.54 0.26 0.27 0.06 0 0 4 0.27 0.06 0 0 0.27 0.13 1.34 0.77 0.27 0.06 0 0 5 2.01 0.19 0 0 3.36 0.64 0.18 0.4 5 0.54 0.19 0 0 6 0.27 0.06 0 0 2.01 0.45 1.68 0.38 0.27 0.06 0.27 0.06 7 0 0 0 0 0.27 0.06 1.61 0.58 0 0 0 0 8 0 0 0 0 0.54 0.13 2.68 0.90 0 0 0 0 9 0 0 0 0 0.67 0.13 7.38 I .09 0.27 0.06 0 0 10 0 0 0 0 0 0 4.03 1.66 0 0 0 0

11 0 0 0 0 2.69 0.70 2.68 0.77 0 0 0 0 12 1.3*+ 0.32 0 0 0.27 0.,06 3.36 0.58 1.61 0.58 0 0 0 0 13 27.52 4.16 0 0 0 0 6.04 2.30 3.23 1.73 1.61 0.38 0.27 0.06 14 3*49 2.37 0.27 0.13 0.27 0.06 5-11 1.28 13.44 3.97 0 0 0.27 0.13 15 0.81 0.38 0 0 0 0 1.88 0.58 1.34 0.58 4.70 0.96 0 0

16 3.49 1.28 0.67 0.13 0 0 1.08 0.58 1.61 0.64 2.01 0.45 0 0 17 0.54 0.13 0.67 0.32 1.34 0. 13 3-49 1.15 0 0 1.08 0.64 0 0

18 1.08 0.32 1.34 0.13 0 0 2.42 1.15 0 0 1.61 0.96 0 0 19 0 0 0 0 0 0 1.88 0.64 0.67 0.06 0 0 0.27 0.06 20 0.67 0.19 0 0 0 0 2.42 0.77 0 0 2.01 0.19 1.34 0.32 21 0.27 0.06 2.01 0.19 0 0 1.34 0.19 0.67 0.06 0 0 3.36 1.41 22 0 0 0.27 0.13 0 0 0 0 0.67 0.26 0 0 0 0 23 0 0 0 0 0 0 0 0 2.96 1.79 3.36 0.77 0.54 0.13 2k 0 0 0 0 0 0 1.34 0.45 0.67 0.06 0.67 0.19 0.27 0.06 25 0 0 0 0 0.27 0.06 0.54 0.19 0 0 2.01 0.32 0 0 26 0 0 0 0 0 0 0 0 2.68 0.64 0.27 0.06 0.27 0.06 27 0 0 0.27 0.06 0 0 0 0 2.68 0.70 0 0 1.61 0.77

28 0 0 0.27 0.06 0 0 2.42 0.83 0.67 0.32 0 0 0 0 29 0 0 0 0 0 0 0.67 0.19 0 0 0 0 0 0 30 0.54 0,11 0 0 0 0 0.81 0.38 0.27 0.06 3.36 0,93 0 0 31 0 0 6*Z1_ 1.09 0,54 0 0

3.. Maximum number of adults/929 cm*" on any trap on a given day.

b. The minimum number of adults/929 cm2 was zero in every case but one* on August 11 this value was 0.8k.

c. Mean number of adult3/929 cm2 from all 6 traps. 150

APPENDIX 19

Number of A. spretulus adults captured per day on 8-vaned sticky tr

Date APR MAY JUN JUL AUG 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4 1 2 3 4

1 0 0 _fi/ _ 3 2 2 3 0 0 0 0 0 0 0 0 0 0 0 2 2 0 0 - - 4 5 0 4 0 0 0 0 0 0 0 0 1 15 0 0 3 1 0 3 6 1 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 4 0 0 1 10 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 5 1 0 4 8 0 0 0 0 0 0 0 0 0 0 0 0 1 6 0 0 6 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 15 0 0 7 110 122 62144 0 0 0 0 0 0 0 0 0 0 0 0 2 3 0 0 S 0 0 0 0 36 26 19 42 0 0 0 0 0 0 0 0 2 1 0 1 9 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 2 0 0 10 157 183 24 33 9 3 4 19 0 0 0 0 0 0 0 0 0 0 0 0 11 0 0 0 0 8 4 4 15 0 0 0 0 0 0 0 0 0 1 0 0 12 3 4 0 0 3 0 0 5 0 0 0 0 0 0 0 0 0 0 0 0 13 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 14 4 2 1 4 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 15 8 8 5 9 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 16 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 17 0 0 0 0 1 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 18 53 28 36 86 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 19 0 0 0 0 0 1 0 1 0 0 0 0 0 3 0 0 0 0 0 0 20 0 0 0 0 2 3 1 2 0 0 0 0 0 1 0 0 0 0 0 0 21 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 22 1 1 1 1 1 0 0 0 0 0 0 0 0 3 0 0 1 2 0 1 23 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 8 15 0 6 24 43 29 6 12 0 0 0 0 0 0 0 0 0 0 0 0 11 16 1 0 25 0 0 0 0 1 1 0 0 1 1 0 0 0 2 0 0 3 17 0 1 26 3 1 5 1 0 0 0 0 0 0 0 0 1 0 0 0 1 13 1 1 27 27 44 14 13 0 0 0 0 0 0 0 0 1 0 0 0 6 39 2 2 28 31 38 5 12 0 0 0 0 0 0 0 0 1 1 0 0 4 10 2 2 29 51 72 11 18 0 0 0 0 0 1 0 0 3 11 0 1 5 30 0 2 30 11 1$ 4 7 0 0 0 0 0 0 0 0 2 5 0 1 4 18 3 4 31 0 0 0 0 0 ? 0 1 2 8 0 0

a. Missing datum. b. Datum preceded by (....) indicates cumulative data for that period of time. 150

APPENDIX 19 s adults captured per day on 8-vaned sticky traps at TPCC during 1978.

MAY JUN JUL AUG SEP OCT 2 3 if 1 2 3 if 1 2 3 J* 1 2 3 if 1 2 3 if 1 2 3 if b/ 2 2 3 0 0 0 0 0 0 0 0 0 0 0 2 • • • • 9 9 9 9 9 • • • 5 0 if 0 0 0 0 0 0 0 0 1 15 0 0 • 9 9 2 i i 5 • • • • 1 0 1 0 0 0 0 0 0 0 0 0 1 0 0 • • • 9 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 14 62 if 10 3 2 0 0 0 0 0 0 0 0 0 0 0 0 0 1 6 0 0 2 5 1 0 0 5 0 0 0 0 0 0 0 0 0 0 0 0 0 3 15 0 0 1 1 I 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 3 0 0 1 7 1 0 0 0 0 0

26 19 if2 0 0 0 0 0 0 0 0 2 1 0 1 • • 9 0 0 0 1 • • • 9 1 1 0 0 0 0 0 0 0 0 0 0 2 0 0 9 • 9 9 2 1 1 3 9 • 9 3 if 19 0 0 0 0 0 0 0 0 0 0 0 0 • • • 2 5 1 6 if if 15 0 0 0 0 0 0 0 0 0 1 0 0 6 13 if 5 0 0 0 0 0 0 5 0 0 0 0 0 0 0 0 0 0 0 0 2 2 3 0 39 66 4 2if 9 9 9 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 if 1 2 2 9 9 9 9 9 9 9 9 9 9 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 6 0 1 9 9 9 9 9 9 « 9 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 *« 9 » 9 9 9 9 9 • • 9 9 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 • 9 • 9 2 3 i 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 If 18 if 12 2 2 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 2 5 1 2 12 9 0 2 1 0 1 0 0 0 0 0 3 0 0 0 0 0 0 2 if 1 2 1 3 1 0 9 9 9 9 3 1 2 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 1 9 9 9 9 9 9 9 • • 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 3 9 9 9 9

0 0 0 0 0 0 0 0 3 0 0 1 2 0 1 • • 9 9 18 17 8 40 • 9 • • t 9 0 0 0 0 0 0 0 0 1 0 0 8 15 0 6 • 9 9 9 0 0 0 0 • 9 • 9 9 9 0 0 0 0 0 0 0 0 0 0 0 11 16 1 0 • 9 9 9 if 2 0 0 1 0 0 1 1 0 0 0 2 0 0 3 17 0 1 0 3 0 i 2 2 0 1 0 0 0 0 0 0 0 1 0 0 0 1 13 1 1 1 0 0 0 0 0 0 2 9 9 9 9 0 0 0 0 0 0 0 1 0 0 0 6 39 2 2 1 1 0 1 9 9 9 9 9 9 9 9 9 0 0 0 0 0 0 0 1 1 0 0 if 10 2 2 0 2 0 0 9 9 9 9 9 • 9 9 9 0 0 0 0 1 0 0 3 11 0 1 5 30 0 2 • 9 9 9 i i i 1 9 9 9 0 0 0 0 0 0 0 2 5 0 1 if 18 3 if 9 9 9 9 6 0 9 0 0 0 0 ? 0 1 2 8 0 0 8 3 0 7

.) indicates cumulative data for that period of time. 151

APPENDIX 20

Number of A. spretulus adults captured per day on 8- vaned sticky traps at KCC during 1978.

APR MAY JUN JUL AUG SEP OCT Date 1 2 1 2 1 2 1 2 1 2 1 2 1 2

0 - 0 0 0 0 0 0 » b/ . • 1 0 0 e » • • 0 - 0 ■ 0 0 1 1 » • 1 2 2 2 1 0 * » • 3 1 0 1 0 0 0 0 0 0 0 • • 0 0 4 1 0 0 0 0 0 0 0 0 0 7 14 1 0 3 0 0 0 0 0 0 0 0 0 0 0 1 0 0 6 0 0 0 0 0 0 0 0 0 0 0 1 0 0 7 136 58 0 0 0 0 0 0 0 0 1 2 0 0 • • 8 0 0 44 147 0 0 0 0 0 1 • • 0 0 « • 9 0 0 2 6 0 0 1 0 1 1 • • 0 1 e • 10 11 40 15 10 0 0 0 0 0 0 • • 2 3 11 0 0 8 16 0 0 0 0 0 0 i i 0 0

12 0 0 3 8 1 0 0 0 0 0 2 l 17 36 * 13 0 2 0 0 0 0 0 0 0 0 0 3 • • s 14 7 3 0 0 0 0 0 0 0 0 2 0 • • • • 15 18 5 0 0 0 0 0 0 0 0 • • • • • 16 0 0 0 0 2 1 0 0 1 0 • • 0 i 17 0 0 3 1 0 0 0 0 0 0 2 6 0 0 18 4 5 1 0 0 0 1 0 0 0 1 5 0 3 19 0 0 1 2 0 0 0 0 0 0 0 2 2 2 • 20 0 0 3 3 0 0 0 0 0 0 2 1 • • 21 0 0 2 0 0 0 2 0 0 0 1 1 t • « 22 0 5 1 1 0 0 0 0 1 1 • • 46 42 • • 23 0 0 0 0 0 0 0 0 0 5 • • 0 0 • • 24 49 19 0 0 0 0 0 0 0 2 • • 5 5 25 0 0 0 0 0 1 0 0 2 5 i 0 0 0 26 0 1 0 0 0 0 1 0 0 2 0 1 2 2 27 28 14 0 1 0 0 0 0 2 3 0 1 0 0 28 39 62 0 0 2 0 0 0 10 6 0 1 0 0 • • 29 44 87 0 1 2 0 0 0 1 3 • • 0 0 30 12 1 0 0 0 0 0 8 3 •• 14 14 0 0 31 0 0 0 0 , 3 7

a. Missing datum. b. Datum preceded by (....) indicates cumulative data for that period of time. 152

APPENDIX 21

Maximum, minimum, and mean numbers of A. spretulus adults captured j at TPCC and KCC during 1978.

JUN JUL Date APR MAY A i n Max Min X Max Min x Max Min X Max*!/ Minb/ V Max3/ Min-

1 - d/ . - 0.38 0 0.21 0 0 0 0 0 0 0.25 0 2 - -- 0.62 0 0.33 0 0 0 0 0 0 1.88 0 3 0.75 0 0.23 0.12 0 0.08 0 0 0 0 0 0 0.12 0 4 1.25 0 0.25 0 0 0 0 0 0 0 0 0 0.12 0 5 1.00 0 0.27 0 0 0 0 0 0 0 0 0 0.75 0 6 0.12 0 0.02 0 0 0 0 0 0 0 0 0 1.88 0 7 18.00 7.25 13.17 0 0 0 0 0 0 0 0 0 0.38 0 8 0 0 0 18.38 2.38 6.54 0 0 0 0 0 0 0.25 0 9 0 0 0 0.75 0 0.23 0 0 0 0.12 0 0.02 0.25 0 10 22.88 1.38 9.33 2.38 0.38 1.25 0 0 0 0 0 0 0 0 o \T\o 11 0 0 0 2.00 * 1.15 0 0 0 0 0 0 0.12 0 12 0.50 0 0.15 1.00 0 o.4o 0.12 0 0.02 0 0 0 0 0 13 0.25 0 0.10 0 0 0 0 0 0 0 0 0 0.12 0 14 0.88 0.12 0.44 0 0 0 0 0 0 0 0 0 0.12 0 15 2.25 0.62 1.10 0 0 0 0 0 0 0 0 0 0.12 0 16 0 0 0 0 0 0 0.25 0 0.06 0 0 0 0.12 0 17 0 0 0 0.38 0 0.15 0 0 0 0 0 0 0.12 0 18 10.75 0.50 4.42 0.12 0 0.02 0 0 0 0.12 0 0.02 0.12 0 19 0 0 0 0.25 0 0.10 0 0 0 0.38 0 0.06 0 0 20 0 0 0 0.38 0.12 0.29 0 0 0 0.12 0 0.02 0 0 21 0 0 0 0,25 0 0.12 0 0 0 0.25 0 o.o4 0 0 22 0.62 0 0.19 0.12 0 0.06 0 0 0 0.38 0 0.06 0.25 0 23 0 0 0 0 0 0 0 0 0 0.12 0 0.02 1.88 0 24 6.12 0.75 3.29 0 0 0 0 0 0 0 0 0 2.00 0 25 0 0 0 0.12 0 0.04 0.12 0 0.06 0.25 0 o.o4 2.12 0 o Q 26 0.62 0 0.23 0 0 0 0.12 0 0.02 0.12 0 • 1.62 0 27 5.50 1.62 2.92 0.12 0 0.02 0 0 0 0.12 0 0.02 4.88 0. 25 o © 28 7.75 0.62 3.90 0 0 0 0.25 0 0.04 0.12 0 1.25 o. 25 29 10.88 1.38 5-90 0.12 0 0.02 0.25 0 0.06 1.38 0 0.31 3.75 0 30 2. 50 0,50 It *6 0.12 0 0.02 0 0 0 0.62 0 0.17 2.25 0.38 31 0 0 0 0 o ro6 1.00 0

a. Maximum number of adults/929 cm on any trap on a given day. b. Minimum number of adults/929 cm on any trap on a given day. c. Mean number of adults/929 cm at all 6 traps. d. Missing datum. 152

APPENDIX 21

m numbers of A. spretulus adults captured per day on 8-vaned sticky traps >78.

AY JUL AUG SEP OCT in X Max Min X Max Min X Max3^ Mi n ^ Max Min X Max Min X 0 0.21 0 0 0 0 0 0 0.25 0 0.04 - d/ - - - -- 0 0.33 0 0 0 0 0 0 1.88 0 0.38 ------0 0.08 0 0 0 0 0 0 0.12 0 0.02 - - - 0.38 0 0.06 0 0 0 0 0 0 0 0 0.12 0 0.02 --- 0.38 0 0.12 0 0 0 0 0 0 0 0 0.75 0 0.15 0.62 0 0.19 0.62 0 0.10 0 0 0 0 0 0 0 0 1.88 0 0.38 0.12 0 0.10 0.12 0 0.02 0 0 0 0 0 0 0 0 0.38 0 0.10 0.88 0 0.25 0 0 0 38 6.54 0 0 0 0 0 0 0.25 0 0.08 - - - 0 0 0 0 0.23 0 0 0 0.12 0 0.02 0.25 0 0.08 --- 0.38 0 0.17 38 1.25 0 0 0 0 0 0 0 0 0 - -- 0.88 0 0.40 50 1.15 0 0 0 0 0 0 0.12 0 0.02 - - - 0 0 0 0 o.4o 0.12 0 0.02 0 0 0 0 0 0 0.38 0 0.21 8.25 0.50 3.88 0 0 0 0 0 0 0 0 0.12 0 0.02 0.50 0 0.25 0.38 0 0.15 0 0 0 0 0 0 0 0 0.12 0 0.02 0.75 0 0.21 0 0 0 0 0 0 0 0 0 0 0 0.12 0 0.02 --- 0 0 0 0 0 0.25 0 0.06 0 0 0 . 0.12 0 0.02 - - - 0 0 0 0 0.15 0 0 0 0 0 0 0.12 0 0.02 - -- 0.25 0 0.10 0 0.02 0 0 0 0.12 0 0.02 0.12 0 0.02 0.62 0.12 0.33 1.50 0 0.54 0 0.10 0 0 0 0.38 0 0.06 0 0 0 0.50 0 0.23 0.38 0 0.19 L2 0.29 0 0 0 0.12 0 0.02 0 0 0 0.25 0 0.10 -- - 0 0.12 0 0 0 0.25 0 0.04 0 0 0 0.38 0 0.12 -- - 0 0.06 0 0 0 0.38 0 0.06 0.25 0 0.12 ------0 0 0 0 0 0.12 0 0.02 1.88 0 0.71 - - - 0 0 0 0 0 0 0 0 0 0 0 2.00 0 0.62 --- 0.62 0 0.33 0 0.0k 0.12 0 0.06 0.25 0 0.04 2.12 0 0.58 - - - 0.25 0 0.10 •d- 0 0 0.12 0 0.02 0.12 0 o o 1.62 0 0.38 0.12 0 0.04 0.25 0 0.12 0 0.02 0 0 0 0.12 0 0.02 4.88 0.25 1.12 0.12 0 0.08 0.12 0 0.02 -3* 0 0 0.25 0 0.04 0.12 0 o O 1.25 0.25 0.71 0.25 0 0.06 0.12 0 0.02 0 0.02 0.25 0 0.06 1.38 0 o.3i 3.75 0 0.85 - - - 0.12 0 0.02 0 0.02 0 0 0 0.62 0 0.17 2.25 0.38 0.83 - - • 1.75 0 0.79 0 0 0,35 0 0.06 1.00 0 0,21 1.00 0 O.SB

p , • cm ' on any trap on a given day. p > cm on any trap on a given day. p i at all 6 traps. 153

APPENDIX 22

Number of A. snretulus adults captured per day in author- made suction black light trap at TPCC during 1977-

Date MAR APR MAY JUN JUL AUG SEP OCT 1 0 0 0 0 1 26 87 2 0 13 0 0 0 4 0 3 0 0 .0 - 1/ 8 1 0 4 0 4 0 - 38 12 0 5 0 8 0 60 13 14 1 6 0 2 0 - 7 10 0 7 0 0 0 26 5 13 0 8 0 0 0 25 23 1 0 9 0 0 0 35 4 77 0 10 2 0 0 20 3 0 0 11 3 0 0 21 4 0 0 12 3 0 0 143 4 2 0 13 8 : s/ 9 8 3 22 0 • 14 1 • 1 97 17 0 0 15 0 34 2 127 10 21 0 16 0 4 1 20 15 60 0 17 0 38 0 38 0 72 0 18 109 40 1 38 0 33 0 19 1 0 6 324 0 30 0 20 5 1 223 0 0 0 21 6 1 63 0 0 1 22 0 8 0 0 5 0 0 23 0 11 3 0 9 199 0 24 0 5 5 0 0 72 0 25 0 7 0 34 0 44 0 26 0 0 2 0 80 1 0 » 27 0 • 0 0 69 1 0 » 28 0 0 • 0 18 47 0 0 29 1 0 • 0 16 6 0 0 30 0 __0 20 __0 24 3 0 31 _JL — 2 111 6 __0

a. Datum preceded by (....) indicates cumulative data for that period'of time. b. Missing datum. APPENDIX 23

Number of A. snretulus adults captured per day in Will-o'-the-Wisp® suction black light trap at LCC during 1977-

Date JU1 AUG SEP OCT

1 13 15 837 2 13 30 0 3 80 21 3 4 31 30 1 5 30 18 4 6 184 19 0 7 130 37 0 8 31 6 0 9 13 33 0 10 114 0 0 11 21 0 0 12 108 0 0 13 42 10 0 14 827 0 0 15 113 60 0 16 38 404 0 17 10 195 0 18 0 113 0 19 242 1 24 0 20 64 11 1 0 21 155 - a/ 0 4 22 73 107 13 0 23 22 225 807 1 24 49 1 9 0 25 90 4 31 0 26 4 55 22 2 27 3 77 2 1 28 13 89 0 0 29 51 17 0 0 30 128 15 -21 0 31 _5i -2k __0

a. Missing datum. 155

APPENDIX 2k

Number of A. spretulus adults captured per day in Will-o'-the-Wlsp® suction black light trap at KCC during 1977.

Date MAY JUNJUL AUGSEP OCT

1 0 1 3 5 3 2 1 1 20 6 0 • 3 * 54 47 0 0 • 4 e 172 132 3 0 5 5 i 12 17 1 1 6 0 0 - 55 0 0 7 0 0 23 70 2 0 8 0 0 6 68 0 0 9 0 0 2 3 18 0 • 10 0 » 3 22 0 2 • 11 0 t• 7 29 1 0 12 0 1 17 12 1 0 13 : s / 1 331 5 0 0 • 14 • 0 126 0 % 319 0 15 l 0 438 3 0 0 16 0 8 97 6 0 0 17 1 0 603 1 5 0 18 0 - 307 0 1 0 19 0 - 632 0 0 0

20 - b/ - 231 0 0 0 21 - 0 45 0 0 0 22 - 0 0 5 0 0 23 17 0 0 2 15 0 24 25 2 0 0 0 0 25 1 0 0 0 2 0 26 0 2 0 15 0 0 • 27 • 0 0 18 0 0 • 28 • 0 44 30 0 • 0 • 29 • 0 53 6 0 0 30 79 __0 516 1 __ 0 0 31 _1S. __ 0

a. Datum preceded by (....) indicates cumulative data for that period of time. b. Missing datum. 156

APPENDIX 25

Mean number of A. spretulus adults captured per day in suction black light traps at TPCC, LCC, and KCC during 1977 ^

Date MAR APR MAY JUN JUL AUG SEP OCT

i 0 0 0 0.50 5-67 15.33 309.00 2 0 13.00 0.50 0.50 11.00 13.33 0 3 0 0 0 54.00 45.00 7.33 1.00 4 0 4,00 0 172.00 67.00 15.00 0.33 5 0 6.50 0 36.00 20.00 11.00 2.00 6 0 1.00 0 - 82.00 9.67 0 7 0 0 0 24.50 68.33 17.33 0 8 0 0 0 15.50 40.67 2.33 0 9 0 0 0 18.50 6.67 42.67 0 10 2.00 0 0 11.50 46.33 0 0.67 11 3.00 0 0 14.00 18.00 0.33 0 12 3.00 0 0 80.00 41.33 1.00 0 13 8 .00 - V 5.00 169.50 16.67 10.67 0 14 1.00 - 0.50 208.00 323.33 0 0 15 0 - 1.00 282.50 42.00 27.00 0 16 0 2.00 4.50 58.50 19.67 154.67 0 17 0 19.50 0 320.50 3.67 90.67 0 18 109.00 20.00 1.00 172.50 0 49.00 0 19 1.00 0 6.00 399.33 0.33 18.00 0 20 5 .00 - 1.00 172.67 3.67 0.33 0 21 6.00 - 0.50 87.67 - 0 1.67 22 0 - 0 24.33 39.00 4.33 0 23 0 14.00 1.50 7.33 78.67 340.33 0.33 24 0 15.00 3.50 16.33 0.33 27.00 0 25 0 4.00 0 41.33 1.33 25.67 0 26 0 0 2.00 1.33 50.00 7.67 0.67 27 0 - 0 1.00 54.67 1.00 0.33 28 0 0 - 0 25.00 55.33 0 0 29 1.00 0 - 0 40.00 9.67 0 0 30 0 0 - 0 222.67 6.33 12.00 0 31 It 00 2. *50 ..6.2,67 16,00 0

a. Calculated for TPCC only from March 28 - May 4, then for TPCC + KCC from May 5 - July 18, and for TPCC + KCC + LCC from July 19 - October 31. b. Missing data from all traps. APPENDIX 26

Number of A. spretulus adults captured per day in Will-o'-the-Wisp® suction "black light traps at TPCC during 1978.

Data MAR APR MAY JUN JUL AUQ SEP OCT 1 2 X 1 2 X 1 2 X 1 2 X 1 2 X 1 2 X 1 2 X 1 2 X 1 0 0 0 0 0 0 8 52 30.0 0 0 0 25 184 104.5 0 0 0 0 1 0.5 2 0 0 0 0 1 0.5 1 12 6.5 0 2 1.0 390 220 305.0 0 0 0 0 0 0 3 0 3 1.5 0 0 0 0 5 2.5 0 2 1.0 48 42 45.0 23 145 84.0 0 0 0 4 0 0 0 0 0 0 0 0 0 0 0 0 1 27 14.0 0 1 0.5 0 0 0 5 0 0 0 0 0 0 0 5 2-5 0 0 0 6 25 15.5 0 11 5.5 0 0 0 6 0 0 0 0 0 0 1 8 4.5 2 30 16.0 39 112 75.5 0 25 12.5 0 0 0 7 1 3 2.0 0 0 0 6 9 7.5 0 3 1.5 3 7 5.0 14 61 37-5 0 0 0 8 0 0 0 11 60 35-5 0 1 0.5 2 9 5.5 8 54 31.0 15 70 42.5 0 0 0 9 0 2 1.0 0 4 2.0 0 0 0 11 73 42.0 29 230 129.5 15 39 27.0 0 0 0 10 0 19 9.5 1 1 1.0 0 1 0.5 0 2 1.0 0 67 33.5 9 37 23.0 0 0 0 11 0 0 0 0 5 2.5 3 16 10.5 0 0 0 9 90 49.5 1 36 18.5 0 0 0 12 0 0 0 0 0 0 0 1 0.5 0 2 1.0 t 28 14.5 28 13 20.5 1 101 51.0 13 0 0 0 0 0 0 0 0 0 64 21 42.5 2 75 38.5 43 85 64.0 1 0 0.5 14 0 0 0 0 0 0 0 0 0 0 0 4 59 31.5 2 88 45.0 13 11 12.0 0 0 0 15 0 0 0 0 0 0 0 0 13 4 8.5 0 3 1.5 1 37 19.0 16 80 48.0 0 0 0 16 0 0 0 0 0 0 0 0 25 78 51.5 1 1 1.0 2 7 4.5 14 67 40.5 0 0 0 17 0 0 0 0 0 0 0 0 4 10 7.0 0 6 3.0 8 19 13.5 18 103 60.5 0 0 0 18 0 0 1 0 0.5 0 1 0.5 0 9 4.5 12 56 34.0 1 106 53.5 42 21 31-5 0 0 0 19 0 0 0 0 0 1 11 6.0 0 0 0 29 140 84.5 6 9 7.5 25 106 65.5 0 0 0 20 0 0 0 0 0 46 42 44.0 1 9 5.0 31 205 118.0 0 1 0.5 17 46 31.5 0 0 0 21 0 0 0 0 0 0 35 31 33.0 0 1 0.5 62 176 119.0 0 0 0 5 7 6.0 0 0 0 22 0 0 0 0 0 0 12 15 13-5 0 0 0 141 264 202.5 1 7 4.0 0 0 0 0 0 0 23 0 0 0 0 0 0 0 6 3.0 0 0 0 52 124 88.0 1 110 55.5 0 0 0 0 0 0 24 0 0 0 0 0 0 0 0 0 0 6 3.0 32 70 51.0 9 95 52.0 0 0 0 0 0 0 25 0 0 0 0 0 0 2 8 5.0 4 8 6.0 23 65 44.0 2 50 26.0 0 0 0 0 0 0 26 0 0 0 0 0 0 2 20 11.0 7 14 10.5 102 184 143.0 1 65 33.0 0 0 0 0 0 0 27 0 0 0 0 0 0 15 100 57.5 4 19 11.5 31 46 38.5 6 149 77-5 0 2 1.0 0 0 0 28 0 0 0 0 0 0 35 101 68.0 3 7 5.0 11 18 14.5 1 30 15.5 0 0 0 0 0 0 29 0 1 0.5 0 0 0 24 66 45.0 0 12 6.0 42 63 52.5 0 26 13.0 0 0 0 0 0 0 30 0 0 0 0 0 0 35 78 56.5 0 9 4,5 28 37 32.5 0 4 2.0 0 0 P 0 0 0 31 , p p . ,.p 8 n 10,5 26 26.0 0 3 1,5 0 0 p

a. Missing datum. 158

APPENDIX 27

Number of A. spretulus adults captured per day Will-o'-the-Wisp® suction black light trap at KCC during 1978.

Date MAR APR MAY JUN JUL AUG SEP OCT

1 0 0 79 0 20 0 0 2 0 0 1 0 3 0 0 3 0 0 0 0 0 1 0 4 0 0 0 0 0 0 0 5 0 0 0 0 0 0 0 6 0 0 1 0 0 1 0 7 0 0 0 0 1 1 0 8 0 1 1 0 12 1 0 9 0 0 0 0 11 0 0 10 0 0 0 0 1 1 0 11 0 0 9 0 1 0 0 12 0 0 0 0 0 0 0 13 0 0 0 0 2 0 0 14 0 0 0 4 4 0 0 15 0 0 0 0 5 0 0 16 0 0 2 0 0 0 0 17 0 0 0 4 0 0 0 18 0 0 0 19 1 0 0 19 0 1 0 21 0 13 0 20 0 0 0 40 0 5 0 21 0 0 0 0 25 0 0 0 22 0 0 0 0 5 0 0 0 23 0 0 0 0 3 16 0 0 24 0 0 0 0 5 5 0 0 25 0 0 0 0 0 4 0 0 26 0 0 1 0 13 0 0 0 27 0 0 3 0 6 0 0 0 28 0 0 8 2 9 0 0 0 29 0 0 141 1 11 0 0 0 30 0 __0 5 __0 9 0 __0 0 31 _0 _li — 1 __0 __0 159

APPENDIX 28

Mean number of A. snretulus adults captured per day in Will-o'-the-Wisp® suction black light traps at TPCC and KCC during 1978.

Date MAR APR MAY JUN JUL AUG SEP OCT

1 C 0 46.33 0 76.33 0 0.33 2 0 0.33 4.67 0.67 204.33 0 0 3 1.00 0 1.67 0.67 29.33 56.33 0 4 0 0 0 0 9-33 0.33 0 5 0 0 1.67 0 10.33 3.67 0 6 0 0 3-33 10.67 50.33 8.67 0 7 1.33 0 5.00 1.00 3.67 25-33 0 8 ‘0 24.00 0.67 3-67 24.67 28.67 0 9 0.67 1.33 0 28.00 90.00 18.00 0 10 6.33 0.67 0.33 0.67 22.67 15.67 0 11 0 1.67 10.00 0 33.33 12.33 0 12 0 0 0.33 0.67 9.67 13.67 34.00 13 0 0 0 28.33 26.33 42.67 0.33 14 0 0 0 0 22.33 31.33 8.00 0 15 0 0 0 5.67 1.00 14.67 32.00 0 16 0 0 0 35-00 0.67 3.00 27.00 0 17 0 0 0 4.67 3-33 9.00 40.33 0 IB 0 0.33 0.33 3.00 29.00 36.00 21.00 0 19 0 0 4.33 0 63-33 5.00 48.00 0 20 0 0 29.33 3.33 92.00 0.33 22.67 0 21 0 0 22.00 0.33 87.67 0 4.00 0 22 0 0 9.00 0 136.67 2.67 0 0 23 0 0 2.00 0 59.67 42.33 0 0 24 0 0 0 2.00 35-67 36.33 0 0 25 0 0 3-33 4.00 29.33 18.67 0 0 26 0 0 7.67 7.00 99.67 22.00 0 0 27 0 0 39.33 7.67 27.67 51.67 0.67 0 28 0 0 48.00 4.00 12.67 10.33 0 0 29 0.33 0 77.00 4.33 38.67 8.67 0 0 30 0 ___0 39.33 l.oo 24.67 1.33 0 0 31 0 _12J3 13.50 0 160

APPENDIX 29

Data tabulated for analysis of variance to test effect of site, month, and sex on survivorship of overwintering adults of A. spretulus.

Observation Site Month Sex Survivorshi p^Mortality Ratio 1 TPCC Sep M 14 0 0.0000 2 TPCC Oct M 105 1 0.0094 3 TPCC Nov M 164 4 0.0238 4 TPCC Dec M 124 5 0.0388 5 TPCC Jan M 158 23 0.1271 6 TPCC Feb M 84 4 0.0455 7 TPCC Mar M 90 4 0.0426 8 TPCC Apr M 1 19 0.9500 9 TPCC May M 0 9 1.0000 10 TPCC Sep F 32 0 0.0000 11 TPCC Oct F 187 1 0.0053 12 TPCC Nov F 276 11 0.0383 13 TPCC Dec F 193 3 0.0153 14 TPCC Jan F 251 23 0.0839 15 TPCC Feb F 109 12 0.0992 16 TPCC Mar F 153 13 0.0783 17 TPCC Apr F 48 ^5 0.4839 18 TPCC May F 0 13 1.0000 19 LCC Sep M 34 0 0.0000 20 LCC Oct M 118 1 0.0084 21 LCC Nov M 67 4 0.0563 22 LCC Dec M 108 2 0.0182 23 LCC Jan M 140 2 0.0141 24 LCC Feb M 151 8 0.0503 25 LCC Mar M 107 11 0.0932 26 LCC Apr M 27 2 0.0690 27 LCC May M 1 7 0.8750 28 LCC Sep F 90 0 0.0000 29 LCC Oct F 250 4 0.0158 30 LCC Nov F 90 10 0.1000 31 LCC Dec F 209 3 0.0142 32 LCC Jan F 267 8 0.0291 33 LCC Feb F 376 26 0.0647 34 LCC Mar F 146 16 0.0986 35 LCC Apr F 114 9 0.0732 36 LCC May F 0 8 1.0000 37 KCC Sep M 17 0 0.0000 38 KCC Oct M 168 6 0.0345 39 KCC Nov M 158 5 0.0307 40 KCC Dec M 164 8 0.0465 41 KCC Jan M 302 10 0.0321 42 KCC Feb M 169 11 0.0611 43 KCC Mar M 163 10 0.0578 44 KCC Apr M 20 42 0.6774 45 KCC May M 0 12 1.0000 46 KCC Sep F 29 0 0.0000 47 KCC Oct F 393 15 0.0368 48 KCC Nov F 286 3 0.0104 49 KCC Dec F 14 0.0488 50 KCC Jan F m 32 0.0653 51 KCC Feb F 320 22 0.0643 52 KCC Mar F 388 29 0.0695 53 KCC Apr F 266 68 0.2036 54 KCC May F 2 23 0.9200 a. Survivorship and mortality calculated as mean of 5 samples taken per site per month. 161

APPENDIX 30

Nutrient mixture for hydroponic culture of turfgrass (after Hoagland and Arnon 1950 and Christians 1977) •

Micronutrient Stock Solutions Constituent g/£ HgO HjBO "boric acid 2.86 MnCl2 *^H20 manganese chloride 1.81 ZnSO^^HgO zinc sulfate 0.22 CuS0^»5H20 copper sulfate 0.08 HgMoO^HgO molybdic acid 0.09 Directions! Add 1 ml of each stock solution for each litre of nutrient solution needed.

Macronutrient Stock Solutions (1 Molar) ml g^cck/ Constituent g/l^ H50 £ nutrient NH|fN03 ammonium nitrate 80.05 2.00 H3PO. phosphoric acid 98.04 1.00 KOH potassium hydroxide 56.11 6,00 CaSO^ calcium sulfate 136.14 ">. 00 M g s c y 7H 2 o magnesium sulfate 246.49 1.00

Iron Stock Solution Constituent g/£ Hg0 Fe2(C^H^0g)-j-HgO iron tartrate 5»00 Directions! Add iron solution at rate of 1 ml/£ of nutrient solution in use, once per week.

l I

162 APPENDIX 31

Artificial diet tested on larvae of A. soretulus (modified from Yamamoto 1969 and Bell and Joachim 1976). Premix Ingredients Wheat germ 90.0 g Casein 63.0 g Sucrose 5^.0 g Brewer's yeast 27.0 g Wesson salt 18.0 g Ascorbic acid 7.2 g Sorbic acid 5.^ g Methyl paraben 1.8 g Cholesterol 1.8 g Streptomycin 0.36 g Other Ingredients Agar 36.0 g Vitamin mixture^/ 9.0 g Aureomycin 0.2 g Formalin (37#) 3.8 ml Poa annua roots and thatch (finely chopped) 75.0 g Directions (1) Place agar into 2-litre flask. Add 1080 ml H20. Heat to boiling while swirling. Remove from heat. (2) Place vitamins and aureomycin into blender. Add 450 ml HgO and blend. Add premix and roots; blend. Add formalin; blend. (3) Pour (1) and (2) into pan and mix thoroughly. Allow to solidify. Cover and refrigerate.

(S) a . Total Vitamin Supplement^ manufactured by the United States Biochemical Corporation, Cleveland, OH.