A STUDY OP SOME ECOLOGICAL FACTORS WHICH AFFECT THE BEHAVIOR OF THE JAPANESE BEETLE IN OHIO DISSERTATION Presented in P a rtia l F ulfillm en t of the Requirements fo r the Degree Doctor of Philosophy in the Graduate School of The Ohio S tate U niversity By RICHARD DEATON WESSEL, B.A., M.Sc. The Ohio S tate U niversity 1951 Approved by?^ Uik &u Dwigmt M. DeLong f / 1 - - ACKNOWLEDGEMENTS To Dr. Joseph B. Polivka, A ssistan t Entom ologist and d irecto r of the Japanese b eetle research at the Ohio A gri c u ltu ra l Experiment S tatio n , fo r h is techn ical d ire c tio n and invaluable assistan ce during the progress of these experim ents and in preparatio n of th is m anuscript, I am deeply Indebted. To Dr. Claude R. Neiswander, Chairman, Department of Entomology a t the Ohio A gricultural Experiment S tatio n , fo r h is valuable advice and au th o rizatio n of th is work; to the various members of the Department of Agronomy of the Ohio A gricultural Experiment S tatio n fo r th e ir suggestions and assistan ce concerning the agronomic aspects of th is problem; and to the Ohio A g ricu ltu ral Experiment S tatio n which made these experim ents p o ssib le, acknowledgement is also rendered. To Dr. Dwight M. DeLong, Professor, Department of Zoology and Entomology a t the Ohio S tate U niversity, fo r h is u n tirin g patience and counsel as my academic advisor; to Dr. D. Ransom W hitney of the Department of M athematics and S ta tis tic s Laboratory of the Ohio S tate U niversity and to Dr. C. R ichard Weaver, A ssistant Entom ologist, a t the Ohio A gricu ltu ral Experiment S tatio n , fo r th e ir assistan ce 2 - - In the evaluation and s ta tis tic a l analyses of these data; and to Clark Robey, Experiment S tatio n photographer, fo r h is aid In the photography concerned w ith th is problem, I also express profound g ra titu d e . To my parents, Dr. and Mrs. C h ristian C. W essel, for th e ir constant In te re st, support, and encouragement esp ecially during my academic years, my h e a rtfe lt apprecia tio n transcends a ll words. To Sue C ailor, who has contributed to th is work in so many charming ways, m ostly unknown to her, I extend my inestim able thanks. 3 - - TABLE OP CONTENTS Acknowledgements................................. ......................................................... 1 In tro d u ctio n ...................................................................................................... 4 The Problem........................................................... ............................................ 5 Review of the L ite ra tu re ....................................................................... 12 F actors of the Edaphic E nvironm ent........................................ 22 Experim ents w ith F e r tiliz e r s ............................................... 22 Summary...................................................................................................... 58 Experim ents w ith Physical F acto rs.................................. 60 Summary..................................................................................................... 70 Experim ents w ith Hydrogen Ion C oncentration (pH)71 Summary...................................................................................................... 8 6 Experim ents w ith Volume W eight and pH in R elation to la rv a l pop u latio n s.................................... 8 8 Summary.................................................................................................... 108 Experim ents w ith Chemical Analyses of S o il 109 Experim ents w ith S o il M oisture.......................................... 112 Summary..................................................................................................... 156 D iscussion of the Problem.................................................................... 159 Summary of the Problem . .................................................................... 161 L ite ra tu re C ited ............................................................................................ 165 Appendix of Supplementary D ata...................................................... 169 The Autobiography........................................................................................ 188 4 - - ' INTRODUCTION The Japanese b eetle (P o p lllla Japonlca Newman) has been present in Ohio since 1931. Each year, since the a rriv a l of th is in sec t, a few new points of in fe sta tio n have been discovered u n til a t present areas of in 1 1 0 fe sta tio n have been recorded, lo cated in 49 of the 8 8 counties of the s ta te . Of the 110 iso la te d in fe sta tio n s, eight have become problem areas. In recent years a tremendous momentum of d isp ersal has developed from these eight p o in ts. The reasons fo r the rap id increase in population in some areas w hile the m ajority of the in fe ste d po in ts re main more or le ss in activ e has been a point of considerable discussion. No im m ediately discernable conditions are apparent to w arrent such behavior when areas of these two types are compared. A pparently there is a com bination of ecological fa c to rs in operation which tend to hold th is in sect in check throughout the m ajority of the in fe sted lo catio n s. To in v estig ate th is behavior and to uncover, if p o ssib le, the complex of ecological fa c to rs responsible fo r th is conduct is the subject of th is d isse rta tio n . 5 - - A STUDY OP SOME ECOLOGICAL FACTORS WHICH AFFECT THE BEHAVIOR OF THE JAPANESE BEETLE IN OHIO The Problem The Japanese b eetle (P o p lllia .japonica Newman) was f ir s t discovered in Ohio in 1931. At th is tim e adult b eetles were trapped in C leveland and in Columbus. At present, years la te r, these areas ex h ib it rem arkably 2 0 d iffe re n t conditions concerning the behavior of th is in s e c t. The C leveland in fe sta tio n has expanded a t an appalling ra te ; u n til at p resen t, 90$ of the area of Cuyahoga and Lake Counties and p a rts of L orain, Medina, Summit, Geauga, and A shtabula Counties compose a large area where the b eetle is continuously d istrib u te d . On the other hand, the Columbus In fe sta tio n is now continuous over less than one- fo u rth of F ranklin County. This difference in area, however, is not the most sig n ific a n t point in the com parison. Throughout Cuyahoga and Lake C ounties there has been a continuous increase in the number of b eetles and accordingly tremendous damage to the tu rf of the many g olf courses, parks, and cem eteries to say nothing of the ravages of the ad u lts upon a ll host p la n ts. The Columbus area, In comparison, has undergone 6 " “ slow expansion and b eetle numbers have never been great enough to cause more than su p e rfic ia l damage or a ttra c t the a tte n tio n of the p u b lic. This is but one of the several sim ilar comparisons th at could be made. C urrently th ere are eig h t problem areas, such as C leveland, in Ohio. These areas in order of prob lem age are (1) Cuyahoga and Lake C ounties, (2) Guernsey County (Cambridge and North Salem), (3) Mahoning and Trumbull Counties (Youngstown and W arren), (4) Licking County (Newark), (5) Columbiana County (East Liverpool and W ellsv ille), ( ) W ashington County (M arietta, e tc .), (7) 6 Jefferso n County (S tu b en v ille), and ( ) Summit County 8 (Akron and B arberton). Figure 1 shows the p resent lo catio n of these areas as w ell as the rem ainder of the in 1 1 0 fe sta tio n s in the sta te , the tim e of discovery of each of these lo catio n s is lis te d in Table 1. The shaded areas around C incinnati and G allap olis show the extent of the b eetle expansion but continuous spraying and so il treatm ent by the U nited S tates Department of A griculture and various sta te agencies have so fa r p re vented the b u ild up of economic populations In these areas. The shaded areas of Wayne, Muskingum, Belmont, and Monroe Counties denote the lo catio n of the b eetle; but, to date, no more than su p e rfic ia l damage has been observed. 7 - - Since the behavior of the Japanese b eetle in Ohio in d icates th a t the in sect has found conditions su itab le in only a few areas, while the m ajority of the lo catio n s have been unsuitable fo r rap id increase in numbers, the facto rs responsible for th is condition should be more c le a rly under stood fo r the purpose of determ ining where the e ffo rts of con trol should be expended and the educational program stressed . Thus fo r the past four years th is author has sought an answer to the question of why th is in sect has become of g reat economic im portance in the problem areas, while other areas, w ith records of the same age, barely m aintain a population, which is freq u en tly im m easureable. The in sect in re la tio n to it s environment, or the study of the in s e c t's ecology, has been the prim ary con sid eratio n . The study of the fa c to rs of the atm ospheric environment such as tem perature, wind, and p re c ip ita tio n have been p a rtia lly sacrificed in favor of working w ith the edaphic fa c to rs. S oil m oisturq pH, f e r til ity , volume w eight, and other physical fa c to rs have received the m ajority of a tte n tio n . 8 - - Table 1 . The location of the Original Japanese Beetle Binds hy Counties and C ities or Townships with the Date of Bind, Humber of Points Within the Area, and the In itia l Humber of Beetles Taken.* Birst Bind Ho . of Ho . 0: City or Trap Be et 1 County Township Dat e Locations Taken Allen Lima I9U3 1 1 Ashland Ashland I9UU 10 23 Londonville I9U5 1 3 Ashtabula Andover 19 50 1 l Ashtabula 1937 7 129 Aust inburg 19 51 - l Cherry Valley 19 51 2 - Conneaut 1935 2 2 Dor set 1951 1 - East Conneaut I9UI 28 91 Geneava I9U5 7 7 Geneva-on-the-Lake 19 51 1 — Jeffer son 1951 - 2 Richmond Center 1951 _ 6 Athens Athens 19^0 1 l H elsonville I9U2 2 2 Belmont Barne sv ille 1939 j■z 3 B ellaire 1937 1 l Bridgeport 1939 1 l Martins Berry 1939 9 13 Butler Hamilton 19^5 l l Middiet own 19 50 1 1 Carroll Carrollton 193S 3 3 Clark Springfield I9H5 l 1 Columbiana Butler Twp 19^2 l 51 East Liverpool 193^ 1 2 Salem 1935 l 1 W ellsville I9H6 1 Several Coshocton Coshocton 1937 15 28 ( Cont inued) ♦These data were compiled from reports on the original in festation s furnished by the State Department of Agriculture, the Bederal Bureau of Entomology and Plant Quarantine, and from the unpublished data of Dr. J. B. Polivka. 9 - - First Find No • of No. 0: City or Trap Beetli County Township Dat e Locations Taken Co sho cton Isleta 19 50 1 k Crawf ord Bucyrus 19^0 3 3 Crestline I9U2 11 20 Cuyahoga Cleveland 1931 1 l Cleveland Heights 1937 1 l East Cleveland 1937 1 1 Er i e Sandusky 193S 1 l Fairf ield Lancast er 1935 2 2 Fayett e Wahsington C.E. 1933 1 1 Franklin Columbus 1931 5 5 Bexley 193S 1 1 Brice 19^9 1 1 G allia G allipoli s 1937 10 12 Geauga Bainbr idge I9U9 1 1 Chardon 19 Hs 2 2 Che sterland 19^9 1 2 Lake Lucerne 1951 b Munson Twp. I9U2 1 b South Eussel 19 51 — b Greene Fairborne I9US 1 2 0 sborne 19 U9 3 b Guernsey Cambridge 1939 2 b Kimbolton 1939 1 1 North Salem 1939 13 69H Hamilton Cincinnati 1937 1 1 Hocking Good Hope Twp. I9U2 1 167 Holme s M iller sburg 19^1 1 1 Huron Bellevue 19^2 67 218 Norwalk iSbo 1 1 Willard 19^3 1 1 Jef f er son Stub enville 1932 H 5 Knox Mount Vernon 19U2 b 5 Lake Madi son 1950 Scout ing Several Mentor 19U1 2 ^7 P ainesville 19^5 20 32 Perry 19U7 2 5 Lawrence Fayette Twp. 19U5 1 1 Ironton 19 51 — 1 Licking Hanover 19 50 1 2 Newark 193s 12 155 ( Continued)